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Full text of "Bulletin of the British Museum (Natural History) Entomology"

56^ 



BULLETIN OF 

THE BRITISH MUSEUM 

(NATURAL HISTORY) 



' G£M£*AL 

3 SEP 19 



ENTOMOLOGY 

Vol. 32 



BRITISH MUSEUM (NATURAL HISTORY) 
LONDON : 1976 



ISSN 0524-6431 



Printed in England by Unwin Brothers Limited 



HtMERAl. ' 

3 5FP 

^ IIWARy a 



CONTENTS 

ENTOMOLOGY VOLUME 32 

No. i. A revision of the African ponerine ant genus Psalidomyrmex Andre 

(Hymenoptera: Formicidae). By B. Bolton i 

No. 2. The butterflies named by J. F. Gmelin (Lepidoptera : Rhopalocera) . 

By R. I. Vane-Wright 17 

No. 3. Revisional notes on African Charaxes, Palla and Euxanthe (Lepidop- 
tera : Nymphalidae). Part X. By V. G. L. Van Someren 65 

No. 4. The species of Chalcidoidea described from North America north of 

Mexico by Francis Walker (Hymenoptera). By B. D. Burks 137 

No. 5. An illustrated list of the type-specimens of the Heliconiinae (Lepidop- 
tera: Nymphalidae) in the British Museum (Natural History). By 
P. R. Ackery & R. L. Smiles 171 

No. 6. A revision of the genus Ptychandra (Lepidoptera : Nymphalidae). By 

H. J. Banks, J. D. Holloway & H. S. Barlow 215 

No. 7. A taxonomic revision of the Tinissinae of the World (Lepidoptera : 

Tineidae). By G. S. Robinson 253 

No. 8. The western Palaearctic Ichneumonidae (Hymenoptera) of British 

authors. By M. G. Fitton 301 



Index to Volume 32 



375 



INDEX TO NOMENCLATURAL CHANGES IN VOLUME 32 



aenea, Callimome 
aethiops, Papilio 
aglidice, Euplaea 
albipes, Eurytoma . 
albipuncta, Tinissa . 275, 

albispiculus, Ephialtes 
alexinus, Brachymeria 
allograptae, Pachyneuron . 
alter, Bathythrix 
amboinensis, Tinissa 269, Pis 
amelanchieris, Syntomaspis 
americana, Haltichella 
americanum, Saranotum 
andricus, Ormyrus . 
anglicanus, Stiboscopus 
anomala, Javra 

approximatus, Exyston cinctulu 
araucariae, Tinissa . 
archippi, Pteromalus 
arctophylax, Hypamblys . 
arridens, Oresbius 
athaliaeperda, Arbelus 
atricilla, Cidaphus 
attenuates, Alexeter 
auriceps, Eurytoma . 
axillaris, Campodorus 

bakeri, Tinissa 
basalis, Eridolius 
bolus, Pseudencyrtus 
braineri, Charaxes 
brevipennis, Agrothereutes 
bridgmani, Promethes 
bruchivorus, Meraporus 
brunneus, Charitopes 
buoliana, Temelucha 
buruensis, Radena . 
bwamba, Palla violinitens 74, 134 



Pis 3 



2, 7 



273. 



(map) 



152 
49 

4i 

163 

8, 12 

340 

164 

154 

336 

11, 12 

151 
165 
147 
153 
34i 
34° 
336 
PI. 12 

*54 
338 
329 
322 

333 
3i6 
162 

353 

PI. 9 
353 
147 
108 

338 
309 
160 

34i 
322 

38 

Pl-5 



38 



caerulans, Ptychandra 

calavius, Notanisomorpha . 

caninae, Otlophorus . 

caproni, Rhorus 

centralis, Palla publius 77, 134 (fig. ] 



chalcites, Tinissa 
chaotica, Tinissa 



274, Pis 3, 8, 



• 239 
. 141 

• 316 

• 317 
, Pis 5. 

6 
12 



105. 



7-', 



Pis 



chintechi, Charaxes . 
chrysopterus, Papilio 
claviger, Danaus juventa . 
collarti, Psalidomyrex procerus 
collinsi, Charaxes usambarae 
Columbiana, Euderus 
confusus, Gelis. 
convoluta, Tinissa 
crassineura, Eurytoma 
cratylus, Neocatolaccus 
croceipes, Phobocampe 
cultellata, Tinissa 
curtisii, Eridolius 
cylindricum, Barylypa 
cyrene, Capellia 
cyrnus, Callitula 

damastes, Achrysocharella 

deficiens, Neostroblia 

deiphon, Trichomalus 

dipsas, Pteromalus 

dobelli, Palla ussheri 71, 134 (map), PI. 3 

dohertyi, Tinissa 279 (fig.), 284, Pis 4, 14 

dymnus, Zatropis .... 157 

elegans, Eridolius .... 353 
elgonae, Euxanthe (Euxanthe) eurinome 

82, 133 (map), 135 (map), PI. 9 



107 

34 

PI. 6 

12 

PI. 19 

144 

3ii 

3. 12 

162 

155 
338 
96, Pis 6, 10 

335 
320 

155 
155 

142 
342 
156 
159 



271, Pis 3, 8 



entellus, Perilampus 
epidius, Aprostocetus 
errantia, Tinissa 
errator, Polytribax . 
eurypon, Trichomalus 
exoticus, Danaus limniace 

fasciatus, Cymodusa 
ficigerae, Semiotellus 
flaccus, Chrysopophagus 
fulvipes, Latibulus . 

glaber, Rhorus. 
glasgowi, Cyrtogaster 
gobryas, Eulophus . 
goliath, Tinissa 
gracilipes, Agrypon . 



• 150 
. 142 

270, Pis 2, 7 

• 339 

• 158 
36, PI. 6 

• 321 

• 159 

• 147 

• 321 



. 3*9 
. 161 

• 145 
287, Pis 5, 15 

• 324 



376 

gracilipes, Alexeter . 
granulatus, Aprostocetus . 

habis, Trichomalus . 
henaultii, Hyposoter 
hephaestos, Jemadia hospita 
hermeas, Zatropis 
heterograpta, Tinissa 
hopei, Aptesis . 
hybreas, Pteromalus. 
hyperboreus, Tymmophorus 



INDEX 



49. 



• 323 
142 

• 158 

• 325 
PI. 6 

■ 159 

• 277 

• 326 
160 

• 337 



illinoensis, Cryptoprymnus . 155 

imbrasus, Neochrysocharis . .145 

impolitus, Homotropus . . . 349 

impressa, Psalidomyrmex feae . 1 1 

indica, Tinissa . . 282, Pis 4, 9, 14 

insularia, Tinissa . . 285, Pis 5, 9, 14 

iphis, Harmolita . . . .163 



jaira, Oreas dubia 
jaira, Papilio . 



39 
39 



kidukaroka, Tinissa 279 (fig.), 287, Pis 5, 15 
kigoma, Palla publius . 77, 134 (map), 

PI. 6 
kigomaensis, Charaxes jahlusa . . 108 

kolae, Lissonota .... 339 
krakatoa, Tinissa . . 267, Pis 2, 11 

kulalae, Charaxes hansali . . .108 

lausus, Acaenacis . . . .159 

leopardus, Danais . . . -37 

leucogyne, Ptychandra 220 (fig.), 221 (fig.), 

246, Pis i, 3, 5 
levana, Stenomesius .... 145 
leytensis, Ptychandra lorquinii 221 (fig.), 

242, PI. 4 
353 
335 
109 
108 

343 

164 

12 

104, PI. 18 



6, 8 



lineola, Eridolius 
linguarius, Hetercola 
lisomboensis, Charaxes imperialis 
loandae, Charaxes 
londinensis, Theroscopus . 
longicornis, Haltichella 
longiscapus, Psalidomyrmex 
lucyae, Charaxes eudoxus . 



mabirensis, Plectroctena mandibularis 
maculicollis, Campodorus . . 353 

madeirensis, Netelia . . . 342 

maderensis, Megastylus . . . 357 

marginatus, Hadrodactylus . .316 



marshalli, Theroscopus . . . 321 

meteori, Eupelminus . . .149 

mindanaensis, Ptychandra lorquinii . 241 



mindorana, Ptychandra 

minio, Pnigalio 
minutus, Lathrolestes 
minutus, Syntactus . 
minyas, Aulogymnus 
miotoni, Charaxes dilutus . 
morleyi, Saotis 
mutina, Danaida limniace 
mysorensis, Tinissa torvella 



221 (fig.), 245, 
Pis 2, 3, 5 

• 145 

• 317 

• 3 10 
146 
109 

■ 34i 

37 

294, Pi. 16 



negrosensis, Ptychandra 250, Pis 2, 3, 5 
nigeriae, Euxanthe (Hypomelaena) trajanus 

89, Pis 15, 16 
nordstromi, Cteniscus lineiger . . 335 



notabilis, Leptacoenites 
notaulius, Stiboscopus 
nox, Tenaris 

nyasicus, Charaxes ethalion 
nyikensis, Charaxes . 

obesus, Psalidomyrmex 
obscurior, Ptychandra lorquinii 
obscurus, Ctenochira 
oculata, Salpinx 
oeax, Mesopolobus . 
ops, Hybophanes 
orontas, Zatropis 



parallela, Tinissa 279 (fig.), 281, Pis 4, 
peregrina, Dusona .... 
peronatus, Campodorus 
phaeus, Charaxes .... 
philippinensis, Tinissa . 277, Pis 
pini, Lamachus .... 

placidus, Hyposoter 
plateni, Ptychandra lorquinii 



241. 



pondoensis, Charaxes 
propinquus, Triptognathus 
protomanica, Charaxes 
proxima, Meloboris . 
proximus, Elachistus 
pseudophaeus, Charaxes 
pythes, Eurytomocharis 

reflexus, Homotropus 
remotus, Entypoma . 
renovatus, Saotis 
romani, Eridolius 



329 

343 

40 

109 

107 

12 
239 
354 

41 
160 

34i 
160 

9, 14 
356 
337 
108 

3. J 3 
336 
326 

(fig-). 

PI. 4 

108 

346 

107 

347 
145 
107 

163 

34i 
338 
34i 
335 



INDEX 



rufescens, Ctenichneumon 


flavocinctus 340 


rufistigma, Pimpla turionellae 


• 344 


rufitibia, Pimpla turionellae 




• 344 


rufulus, Tryphon 






• 354 


rufus, Platyrhabdus . 






• 339 


rusticus, Xorides 






• 33° 


sallyae, Psalidomyrmex 




•9, 15 (fig-) 


scutellatus, Mesoleptus 




• 316 


senegala, Charaxes fulvescens 




103 


sericans, Astiphromma 






323 


scoticus, Aptesis 






339 


solus, Encyrtus 






148 


spiracularis, Pimpla alternans 




343 


splenium, Astiphromma 






323 


strobiloides, Callimome 






!5i 


subfasciatus, Rhorus 






354 


subrufa, Polysphincta 






319 


suna, Charaxes anticlea 






108 


suturalis, Apophua . 






340 


syrphi, Spalangia 






154 


talboti, Ptychandra • 220 


(fig.) 


, 248, Pis 1, 








3. 5 



teredon, Tenuipetiolus . 
thomsoni, Theroscopus 
tibialis, Bassus 
timaea, Zatropis 
tomentosus, Ctenopelma 
transversella, Tinissa 
tricolor, Homotropus 
trypherus, Cyrtogaster 

ulixes, Pyrrhopyga . 
unicinctus, Polyblastus 
ustulatus, Xenoschesis 

vansoni, Charaxes 
variata, Charaxes 
vectius, Psyllaephagus 
ventralis, Hadrodactylus 



377 

• 163 

• 336 

• 309 
. 161 

326 
283, Pis 4, 14 

• 349 

. 161 



50 
319 
33° 

108 
108 
148 
323 



whitei, Charaxes cithaeron cithaeron . 105, 

Pi. 19 



xylonomoides, Acrolyta 



339 



\J\ I i 



... 



1 8 APR m 

A REVISION U 

OF THE AFRICAN PONERINE ANT 
GENUS PSALIDOMYRMEX ANDRE 
(HYMENOPTERA : FORMICIDAE) 



B. BOLTON 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 1 

LONDON : 1975 



A REVISION OF THE AFRICAN 

PONERINE ANT GENUS PSALIDOMYRMEX 

ANDRE (HYMENOPTERA : FORMICIDAE) 



BY 

BARRY BOLTON 



Pp. i-i6, 6 Text-figures 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 1 

LONDON : 1975 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 1 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.) 



ISSN 0007-1501 



Trustees of the British Museum (Natural History), 1975 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 10 April, 1975 Price £1-30 



A REVISION OF THE AFRICAN 

PONERINE ANT GENUS PSALIDOMYRMEX 

ANDRE (HYMENOPTERA : FORMICIDAE) 

By B. BOLTON 



CONTENTS 



Synopsis 

Introduction 

Abbreviations of museums 

Measurement and indices 

Definition of the genus 

List of species 

Key to species (Workers) 

The foveolatus-group 

The procerus-group 
Acknowledgements 
References . 
Index 



Page 

3 
3 
4 
4 
5 
6 
6 

7 
ii 

"4 

14 
16 



SYNOPSIS 

The ant genus Psalidomyrmex of the Ethiopian Region s revised. Keys to species and descrip- 
tions are presented. Six species are recognised, one of which is described as new, and five new 
synonyms are established. 



INTRODUCTION 

Psalidomyrmex is a small, well-defined genus of ponerine ants containing only six 
species. It is peculiar to the wet forest zones of sub-Saharan Africa, particularly 
to the rain forest belts of West and Central Africa although some species do occur 
also in the Ugandan forests and one species is found on the offshore Principe Island, 
in the Gulf of Guinea. 

Nests are constructed in rotten wood, usually in an advanced state of decay, 
or more rarely directly into the soil beneath a log. Individual foraging workers 
are found below rotten logs and in deep leaf litter and log mould. The food of 
the species is not known but the specialized mandibles of both species-groups of 
the genus imply a specialized diet. 



4 B. BOLTON 

Apart from scattered descriptions of new forms the only previous study of the 
genus was presented by Wheeler (1922) who gave a distribution map of the genus 
and a key to the then-known species. In many ways this key is unsatisfactory 
and in fact breaks down on the first couplet, even when only the species included 
in it are considered, as foveolatus is more closely related to reichenspergeri and shares 
the same mandibular configuration. In his key Wheeler implies that foveolatus 
has a mandibular structure similar to that of procerus, which is not the case, and 
further states that in foveolatus the scapes are 'reaching or surpassing the posterior 
corners of the head', which they do not. 

The present paper recognizes two species-groups within Psalidomyrmex, each 
containing three species, based primarily upon the structure of the mandibles. 
The groups and their constituent species are discussed below. 



ABBREVIATIONS OF MUSEUMS 

AMNH, New York American Museum of Natural History, New York, U.S.A. 

BMNH British Museum (Natural History), London, U.K. 

IE, Bologna Istituto di Entomolgia del'Universita, Bologna, Italy. 

MCZ, Cambridge Museum of Comparative Zoology, Cambridge, Mass., U.S.A. 

MNHN, Paris Museum National d'Histoire Naturelle, Paris, France. 

MRAC, Tervuren Musee Royal de l'Afrique Centrale, Tervuren, Belgium. 

NM, Basle Naturhistorisches Museum, Basle, Switzerland. 

NM, Bulawayo National Museum, Bulawayo, Rhodesia. 

MEASUREMENTS AND INDICES 

Total Length (TL). Total outstretched length of the individual, from mandibular 
apex to gastral apex. 

Head Length (HL). The straight-line length of the head in perfect full-face view, 
measured from the mid-point of the anterior clypeal margin (excluding the 
prominent labral lobe) to the posteriormost point of the occipital margin. (In 
species with a concave occipital margin the head length is measured to the mid- 
point of a line connecting the posterolateral projections.) 

Head Width (HW). The maximum width of the head measured behind the eyes 
in full-face view. 

r 1, v t a in\ HW x I0 ° 
Cephalic Index (CI). — 

HL 

Scape Length (SL). The straight-line length of the antennal scape excluding 
the basal constriction or neck. 

Scape Index (SI). — — — 

HW 

Pronotal Width (PW). The maximum width of the pronotum in dorsal view. 

Petiole Length (PL). The length of the petiole in profile from the anterior process 
to the posteriormost point of the tergite, where it surrounds the gastral articula- 
tion. 



REVISION OF GENUS PSALIDOMYRMEX 5 

Dorsal Petiole Width (DPW). The maximum width of the petiole in dorsal view. 

Dorsal Petiole Index (DPI). DPW X I0 ° 

v ; PL 

All measurements are expressed in millimetres. 



DEFINITION OF THE GENUS 

PSALIDOMYRMEX Andre 

Psalidomyrmex Andre, 1890 : 313. Type-species: Psalidomyrmex foveolatus Andre, 1890 : 314; 
by monotypy. 

Worker. Black or red-brown ants belonging to the tribe Ponerini. Monomorphic but with 
notable size variation in some species. Lifeway cryptic, with nest sites usually in rotten wood. 
Size ranges from medium to large, TL ca 9-0— 16-0. 

Mandibles either elongate and falcate, with a concave apical (masticatory) margin which 
is equipped with a number of short teeth near the basal angle (foveolatus-group, Text-fig. 1), 
or the mandibles subtriangular with an elongate apical tooth, the apical margin more or less 
straight, edentate (procerus-group, Text-fig. 2). In both groups the basal angle is rounded, 
not acute, and a broad mandibular groove is present which commences laterodorsally and runs 
down the outer margin of each mandibular blade; the blades cross over at rest. Labrum 
distinct, prominent, in dorsal view projecting beyond the anterior margin of the clypeus as a 
rounded, usually transversely striate lobe. Clypeus short, transverse. Palp formula of 
maxillary 3, labial 4 segments (dissections of foveolatus, procerus, reichenspergeri, wheeleri). 
Lateral lobes of frontal carinae strongly expanded, completely concealing the antennal 
insertions. Antennae with 12 segments, the second funicular segment usually noticeably 
longer than the third. Eyes present, usually of moderate size but reduced in some species. 
Dorsal alitrunk with promesonotal suture distinct, metanotal groove reduced or virtually 
absent. Propodeal dorsum usually with a median, longitudinal, narrow groove or impression 
which in some species is also present on the posterior portion of the pronotum. Middle and 
hind tibiae each with a single pectinate spur, without a smaller, lateral spur. Pretarsal 
claws simple. Petiole nodiform. Gaster strongly constricted between the first and second 
segments. 

Female. As worker but alate, the alitrunk with a full complement of flight sclerites. Ocelli 
present. 

Male. Mandibles very reduced, edentate, short, roughly rectangular in shape and failing 
to meet apically at full closure. Antennae with 13 segments, filiform, the scape and first 
funicular segment short, their combined length less than that of the second funicular segment. 
Eyes large, ocelli present. Lobes of frontal cairnae small, raised, only partially covering the 
antennal insertions. Alitrunk with a full complement of sutures and flight sclerites. Pro- 
notum strongly developed, not overhung by the mesoscutum in profile. Mesocutum without 
notauli, parapsidal furrows present but usually masked by the sculpturation. Scutellum 
swollen, usually somewhat dome-shaped in profile and with a dorsal, longitudinal impression. 
Hind wings with anal lobe present. Middle and hind tibiae each with a single pectinate spur, 
lateral spurs absent. Pretarsal claws with a tooth at or distal to the midlength. Gaster 
strongly constricted between first and second segments. Pygidium blunt or truncated apically, 
not produced into a curved spine; hypopygium broadly Y-shaped or bluntly truncated 
apically with curved lateral angles. Genitalia rectractile. Gonopalpi present, usually concealed 
by the pygidium. 



6 B. BOLTON 

The genus Psalidomyrmex is most closely related to Plectroctena F. Smith, another 
genus which is confined to the Ethiopian Region. A discussion of the relationships 
and derivation of these genera has been presented in a recent revision of Plectroctena 
(Bolton, 1974). 

LIST OF SPECIES 
Jo veolatus-group 
foveolatus Andre 
reichenspergeri Santschi 

mabirensis (Arnold) syn. n. 
sally ae sp. n. 
procerus-group 
jeae Menozzi 

feae var. impressa Menozzi syn. n. 
procerus Emery 

longiscapus Santschi syn. n. 
obesus Wheeler syn. n. 
procerus st. collarti Santschi syn. n. 
wheeler i Santschi 

A further species, clavicornis Bernard (1952 : 209), has been removed from the 
genus and transferred to Bothroponera as a synonym of B. talpa (Andre) (W. L. 
Brown, personal communication of paper in preparation). I have examined 
the holotype female of clavicornis (in MNHN, Paris) and concur with Professor 
Brown. 

KEY TO SPECIES 

Workers 

1 Mandibles falcate, the masticatory margin concave and with a number of short 

teeth, at least near the basal angle (Text-figs 1,5). . . . • • 2 

- Mandibles subtriangular with an elongate apical tooth ; the masticatory margin eden- 

tate and straight or nearly so (Text-figs 2, 6) . . . . . . . 4 

2 In dorsal view the first gastral tergite strongly narrowed anteriorly (Text-fig 3). 

Dorsal surface of first gastral tergite with small punctures, the diameters of which 
are less than the distances separating them. Petiole longer than broad, DPK. 
100. Large species HW> 2 -50, PW> 1.70. Full adult colour black. (Uganda, 
Cameroun, Zaire) ....... reichenspergeri (p. 8) 

- In dorsal view the first gastral tergite not narrowed anteriorly (Text-fig 4) . Dorsal 

surface of first gastral tergite coarsly foveolate, the diameters of the foveolae equal 
to or greater than the distances separating them. Petiole as broad as or broader 
than long, DPI 100 or more. Smaller species, HW < 2-50, PW <.i-jo. Full 
adult colour red-brown .......... 3 

3 Spaces between foveolae on pronotal dorsum and first gastral tergite densely striate. 

Antennal scapes relatively short, 5/ < 75. Eyes small, maximum diameter 

< 0-26. (Sierra Leone, Ivory Coast, Ghana, Nigeria) . . foveolatus (p. 7) 

- Spaces between foveolae on pronotum and first gastral tergite unsculptured, mostly 

smooth and shining. Antennal scapes relatively long, SI > 80. Eyes larger, 
maximum diameter > 0-26. (Ghana) ..... sallyae (p. 9) 

4 Antennal scapes relatively long, 5/ 90 or more (range 90-102) ; head relatively 

narrow, CI 89 or less (range 84-89) ; petiole longer than broad, DPI < 100. (Cam- 
eroun, Zaire) .......... wheeleri (p. 13) 



REVISION OF GENUS PSALIDOMYRMEX 7 

Antennal scapes shorter, SI 88 or less (range 79-88); head relatively broad, CI 91 

or more (range 91-96); petiole broader than long, DPI ~> 100 . . . . 5 

Expanded lobes of frontal carinae smooth and shining over the site of the antennal 
insertions, unsculptured apart from a few striae centrally and basally. Median 
portion of clypeus unsculptured or with transverse striae. (Principe I.) . feae (p. 1 r) 

Expanded lobes of frontal carinae uniformly (but sometimes faintly) striate over the 
site of the antennal insertions. Median portion of clypeus longitudinally striate. 
(Ghana, Uganda, Cameroun, Gabon, Zaire) ..... procerus (p. 12) 



The FOVEOLATUS-Grovp 

Characterized by the mandibles which are elongate-falcate and dentate in part. 

In the three species constituting this group the mandible has apparently been 
modified from a basically subtriangular shape into a falciform structure. This 
has been achieved by the elongation of the apical margin distal to the rounded 
basal-internal angle. The apical (masticatory) margin has become concave and 
runs without interruption into the long, curved apical tooth so that the two are in 
effect a single structure. The apical margin retains short teeth in the part of its 
length closest to the basal angle, and in some specimens small teeth occur upon 
the curve of the basal angle itself. The basal portion of the mandible is also some- 
what elongated (Text-figs 1, 5). 

Of the three species known with mandibles as defined above, two (foveolatus 
and sallyae) are closely related, relatively small red-brown species which appear 
to be restricted to West Africa. The former has a wide range, from Sierra Leone 
to Nigeria, but the latter is as yet only known from two collections made in Ghana. 

The third species, reichenspergeri, also has a wide range, but interestingly has 
not yet been found in any territory in which foveolatus is known to occur. The 
reverse also applies as foveolatus is not reported from Cameroun, Zaire or Uganda, 
the known range of reichenspergeri. This last species is larger than its West African 
counterparts and is black in colour. 



Psalidomyrmex foveolatus Andre 
(Text-fig. 4) 

Psalidomyrmex foveolatus Andre, 1890:314. Syntype workers, Sierra Leone (A. Mocquerys) 
(MNHN, Paris) [examined]. 

Diagnosis of worker. Mandibles falcate. Antennal scapes with SI < 75. First gastral 
tergite not narrowed anteriorly; the foveolae on this segment and the pronotal dorsum with a 
dense striation between them. DPI 100 or more. 

Further description. Worker. TL 9-4-10-2, HL 1-94-2-12, HW 1-68-1-84, CI 84-87, 
SL 1-20-1-32, SI 66-74, PW 1-24-1-32, PL 0-84-0-88, DPW 0-84-0-92, DPI 100-105 ( IO 
measured). 

Mandibles falcate, their apical margins with a number of small teeth distal of the basal 
angle. In some specimens the teeth are acute but in others low and rounded; occasionally 
teeth occur on the curvature of the basal angle itself. Eyes small, maximum diameter 
ca 0-16-0-22, distinctly less than the maximum width of the scape. Clypeal suture effaced, 
very reduced, usually not breaking the sculpture ; not visible in more deeply coloured individuals. 



8 B. BOLTON 

General outline of head similar to that of sallyae (Text-fig. 5). Promesonotal suture distinct 
and impressed. Metanotal groove absent, in profile not impressed, the mesonotum and propo- 
deum forming a continuous convexity. In dorsal view the original track of the metanotal 
groove is visible in some specimens but usually it cannot be discerned, and the dorsal longitudinal 
sculpture is always unbroken. Propodeal dorsum with a broad, median longitudinal impression 
or groove, at least posteriorly. The groove is generally broadest posteriorly, narrowing anteriorly. 
Pronotal dorsum without such a groove. Petiole in dorsal view usually slightly broader 
than long, with a rounded anterior and transverse posterior face. Dorsal surfaces of body 
and head with erect hairs, the appendages with hairs on all surfaces. 

Sculpture everywhere basically of a dense, longitudinal striation with scattered foveolae. 
Striate sculpturation is also present on the mandibles, antennal scapes and legs. The expanded 
lobes of the frontal carinae are for the most part smooth, with striae restricted to the basal and 
internal portions. On the sides of the alitrunk the striation is finer than on the dorsum, and 
on the sides of the propodeum may be very fine indeed. Sides of petiole foveolate, the spaces 
between the foveolae reticulate-punctate. Dorsum of petiole as sides but the sculpture between 
foveolae much less intense than on the sides, superficial, in places absent on some specimens. 
Full adult colour a deep red-brown. 

Female. As worker but with ocelli, and the alitrunk with flight sclentes. TL 10-4-11-6, 
HL 2-08-2-20, HW 1-80-1-88, CI 85-86, SL 1-28-1-40, SI 71-74, PW 1 -48-1 -76, PL 0-92-1-00, 
DPW 0-96-1 -04, DPI 104. Maximum diameter of eye 0-30-0-36 (3 measured). 

P. foveolatus is the commonest and most widely distributed species of the genus 
in the West African forest zone, ranging from Sierra Leone to western Nigeria. 
Nests are made in rotten logs, usually in an advanced state of decay, and workers 
forage singly either in the log or in the surrounding leaf litter. 

P. sallyae, described below, is certainly the species most closely related to foveo- 
latus, but the differences separating the two are numerous and are tabulated under 
sallyae. P. foveolatus itself appears to show but little variation over its wide range. 

Material examined. 

Sierra Leone: no date (ex coll. F. Smith). Ivory Coast: Lamto, Toumodi 
(/. Le'vieux). Ghana: Tafo (B. Bolton); Tafo, numerous series (D. Leston); Legon 
(D. Leston); Kibi (D. Leston); Wiawso (D. Leston); Asamankese (D. Leston). 
Nigeria: Ibadan (Booker). 



Psalidomyrmex reichenspergeri Santschi 
(Text-figs 1, 3) 

Psalidomyrmex reichenspergeri Santschi, 1913 : 302. Holotype worker, Cameroun: Molunda 

(A. Reichensperger) (NM, Basle) [examined]. 
Plectroctena mandibularis subsp. mabirensis Arnold, 1954 : 293 figs 3, 3a. Syntype workers, 

Uganda: Mabira Forest, 21.V.1952 (G. Arnold) (NM, Bulawayo) [examined]. Syn. n. 
Psalidomyrmex mabirensis (Arnold); Bolton, 1974 : 334. 

Diagnosis of worker. Mandibles falcate. Antennal scapes with S7> 75. First gastral 
tergite narrowed anteriorly, with small punctures but without foveolae. Large species, 
HW > 2-50. DPI < 100. 

Further description. Worker. TL 14-8-15-8, HL 3-28-3-48, HW 2-80-3-16, CI 85-90, 
SL 2-20-2-44, SI 76-78, PW 1-80-2-20, PL 1-36-1-48, DPW 1-12-1-32, DPI 82-90 (5 measured). 

Mandibles falcate and with a number of short teeth on the masticatory margin close to the 
basal angle. In some specimens the teeth continue onto the curvature of the basal angle. 



REVISION OF GENUS PS A LI DOM Y RM EX 9 

Sides of head feebly convex, the occipital margin broadly but shallowly concave. Eyes with 
maximum diameter ca o-32-o-42, slightly less than the maximum width of the antennal scape. 
Promesonotal suture strongly impressed. Metanotal groove distinct in dorsal view, impressed 
in profile and separating the mesonotal convexity from that of the propodeum. Propodeal 
dorsum with a median, longitudinal groove running from the mid-point of the metanotal groove 
to the declivity. Petiole in dorsal view slightly longer than broad. In dorsal view the first 
gastral tergite strongly narrowed anteriorly so the extreme anterior surface is narrower than 
the posterior face of the petiole node. Erect hairs absent from the mesonotum and propodeum 
and usually absent from the first gastral tergite although on this surface a few may be present 
posteriorly. 

Dorsum of head coarsely punctate, the spaces between the coarse punctures strongly longi- 
tudinally striate. Dorsal alitrunk with scattered coarse punctures, less dense than on the head 
and tending to be smaller and more numerous on the propodeum than the pronotal dorsum. 
Spaces between punctures on the pronotal dorsum tend to be smooth and shining although a 
few striae may be present, especially posteriorly. Mesonotal dorsum generally with weak 
striation; the propodeal dorsum finely but distinctly striate between the punctures. Sides of 
alitrunk with scattered punctures and fine dense striation. First gastral tergite with scattered 
punctures which are smaller than those on the head, the spaces between them mostly smooth 
but sometimes with very faint, superficial striation. Second gastral tergite as first but the 
striation much more distinct. Scapes and lobes of frontal carinae not striate. Full adult colour 
black. 

Male. A specimen labelled as the male of reichenspergeri (det. Santschi) is present in the 
Santschi collection (NM, Basle). It bears the data 'Kamerunberg, Soppo 730 m 1912 (v. 
Rothkirch).' 

This specimen differs from others in the genus which I have examined by having a narrow, 
very broadly Y-shaped subgenital plate, the dorsal arms of which are long and strongly curved. 
In other males examined, which I attribute to procerus, the subgenital plate is broad with short, 
thick, feebly recurved free angles. 

The form of the mandibles relates this species to foveolatus and sallyae but reichen- 
spergeri is larger than both these species and is black. The main characters separ- 
ating it from its relatives are given in the key to species and the diagnoses. 

Arnold (1954) originally described mabirensis as a subspecies of Plectroctena 
mandibularis Smith, but his figures and description were sufficient to indicate 
that mabirensis should correctly be placed in Psalidomyrmex, where it was later 
transferred (Bolton, 1974). Examination of the syntypes of mabirensis and direct 
comparison of them with the holotype of reichenspergeri now shows that the two 
names are synonyms. 

Material examined. 

Cameroun: no. 1343 (G. Terr on). Zaire: Akenge (Lang & Chapin). 

Psalidomyrmex sallyae sp. n. 

(Text-fig. 5) 

Diagnosis of worker. Mandibles falcate. Antennal scapes with SI > 75. First gastral 
tergite not narrowed anteriorly; the foveolae on this segment and the pronotal dorsum without 
dense striation between them. DPI > 100. 

Further description. Holotype worker. TL io-o, HL 2-08, HIV 1-76, CI 85, SL 1-48, 
SI 84, PW 1-36, PL o-88. DPW o-Q2, DPI 104. 



io B. BOLTON 

Outline shape of head as shown in Text-fig. 5. Mandibles falcate and with a number of 
short teeth on the apical margin which extend onto the rounded basal angle as low crenulations. 
Eyes with maximum diameter ca 0-30, about equal to the maximum width of the scape. 
Clypeal suture effaced, not breaking the sculpturation. Alitrunk in profile with the promeso- 
notal suture strongly impressed, the metanotal groove more weakly so but sufficiently marked 
to separate the mesonotal curvature from that of the propodeum. In dorsal view the metanotal 
groove poorly developed but easily discernible. Propodeal dorsum with a median longitudinal 
groove running from the metanotal groove to the declivity. Pronotum without such a median 
groove. Dorsal surfaces of head and body with numerous erect hairs, which are also present 
upon the appendages. 

Dorsal and lateral surfaces of head, alitrunk, petiole and gaster foveolate. On the dorsum 
of the head the spaces between foveolae are faintly longitudinally striate, the striation weak, 
virtually effaced in places, stronger on the sides of the head below and behind the eyes. 
Pronotal dorsum without striae between the foveolae although on the mesonotum and 
propodeum one or two weak striae may be present on the otherwise smooth surfaces. Metanotal 
groove with a band of short striae which project for a short distance both anteriorly and 
posteriorly. Sides of alitrunk with spaces between foveolae striate. Sides and dorsum of 
petiole with fine punctures or superficial reticulation between the foveolae. Gaster with spaces 
between foveolae unsculptured, both dorsally and ventrally. Mandibles striate; lobes of 
frontal carinae unsculptured. Full adult colour a deep red-brown. 

Paratype workers. As holotype but one a teneral, with head and alitrunk orange-brown, 
the gaster darker. Dimensions of paratypes: TL 9-6-11-4, HL 2-00-2-32, HW 1-68-2-00, 
CI 83-86, SL 1-40-1-60, 57 83-85, PW 1-32-1-56, PL 0-84-1-00, DPW 0-92-1-08, DPI 104-109. 
Maximum diameter of eye ca 0-28-0-34 (4 measured). 



Holotype worker, Ghana: Tafo, 23.vii.1966, ant ecology sample 120 (D. Leston) 
(BMNH). 

Paratypes. Ghana: 3 workers, same data as holotype, one of them also bearing 
the number 806; 1 worker from the same locality but 19.vii.1966, ant ecology sample 
no (D. Leston) (BMNH; MCZ, Cambridge; AMNH, New York; NM, Bulawayo). 

This species is very closely related to foveolatus and occurs within the range of 
that species. The following table illustrates the main characters serving to differ- 
entiate workers of the two species. 



foveolatus 



sallyae 



Antennal scapes shorter, 5/ 66-74. 
Maximum diameter of eye 0-16-0-22, 
less than maximum width of scape. 

Metanotal groove vestigial or absent, 
not impressed in profile. 
Pronotum and first gastral tergite with 
strong striation between foveolae. 



Antennal scapes longer, SI 83-85. 

Maximum diameter of eye 0-28-0-34, 

about equal to maximum width of 

scape. 

Metanotal groove visible in dorsal view, 

impressed in profile. 

Pronotum and more especially first 

gastral tergite without striae between 

foveolae. 



revision of genus psalidomyrmex n 

The PROCERUS-Grovp 

Characterized by the mandibles which are subtriangular with an elongate apical 
tooth, and with the apical (masticatory) margin edentate. 

In this group the mandibles have retained a rather more generalised shape than 
is encountered in the previous group. The mandibles in procerus and its allies 
are subtriangular, with a long, curved apical tooth which is very broad and appears 
to be truly a continuation of the mandibular blade. The apical margin proximal 
to this tooth is more or less straight in full-face view and is edentate, with a fine 
and quite narrow cutting edge. The basal angle is broadly rounded, and the 
external margin is somewhat concave at about its midlength. The basal portion 
of the mandible is not markedly elongate (Text-figs 2, 6). 

The three species placed in the procerus-gvoup are very closely related, and 
procerus itself is the most variable species yet known in the genus in terms of size, 
sculpture and hairiness. The least known species, feae, has at present only been 
reported from Principe Island in the Gulf of Guinea, but of the others wheeleri 
is known from Cameroun and Zaire whilst procerus is widely distributed in West 
and Central Africa and also occurs in Uganda. 



Psalidomyrmex feae Menozzi 
(Text-fig. 6) 

Psalidomyrmex feae Menozzi, 1922 1349. Syntype workers, female, male, Principe I.: Roca 

Infante Don Henrique, iii.1901, 100-300 m (L. Fea) (IE, Bologna; MCZ, Cambridge) 

[examined] . 
Psalidomyrmex feae var. impressa Menozzi, 1922 : 352. Syntype workers, male, Principe I.: 

Roca Infante Don Henrique, ii.1901, 200-300 m (L. Fea) (IE, Bologna; MCZ, Cambridge) 

[examined]. Syn. n. 

Diagnosis of worker. Mandibles subtriangular. Expanded lobes of frontal carinae 
over antennal insertions not striate. SI < 90, DPI > 100. 

Further description. Worker. TL 10-8-12-2, HL 2-08-2-36, HW 1-96-2-24, CI 92-95, 
SL 1-56-1-80, SI 79-81, PW 1-52-1-68, PL 1-00-1-08, DPW i-i2-i-2o, DPI 108-116 (4 
measured). 

Very closely related to procerus and separated from it only by the following. 

1. Expanded lobes of frontal carinae smooth and shining in feae, striate in all specimens of 

procerus examined. 

2. Median portion of clypeus immediately in front of the lobes of the frontal carinae 

unsculptured or with feeble transverse striae in feae, longitudinally striate in procerus. 

3. Size range in feae at lower end of procerus range. 

As only the type-series of workers and a single queen of feae and its absolute 
synonym impressa have been examined the consistency of the characters listed 
above cannot be guessed. However, in the material examined they serve to separate 
the two species and it has been decided to retain feae as a distinct species until 
further material becomes available. 



12 B. BOLTON 

Psalidomyrmex procerus Emery 
(Text-fig. 2) 

Psalidomyrmex procerus Emery, 1901 : 50. Syntype females, male, worker, Cameroun (Conradt) 

(NM, Basle) [examined]. 
Psalidomyrmex longiscapus Santschi, 1920 : 8. Holotype female, Gabon: Samkita (F. Faure) 

(NM, Basle) [examined] . Syn. n. 
Psalidomyrmex obesus Wheeler, 1922 : 92, fig. 19. Syntype workers, Zaire: Medje (Lang 6- 

Chapin) (AMNH, New York; MCZ, Cambridge) [examined]. Syn. n. 
Psalidomyrmex procerus st. collarti Santschi, 1937 : 74. Holotype worker, Zaire: Ituri, 

Matenda, 22. ix. 1929 (A. Collart) (MRAC, Tervuren) [examined]. Syn. n. 

Diagnosis of worker. Mandibles subtriangular. Expanded lobes of frontal carinae 
striate. 57 < 90. DPI > 100. 

Further description. Worker. TL 11 -4-16-2, HL 2-28-3-20, HW 2-12-3-04, CI 91-96, 
SL 1-76-2-48, SI 80-88, PW 1-72-2-36, PL 1-08-1-56, DPW 1-24-1-68, DPI 106-118 (20 
measured). 

Mandibles elongate-subtriangular, edentate (Text-fig. 2). Sides of head convex, the occipital 
margin impressed medially, the outline shape generally as in Text -fig. 6. Eyes of moderate 
size, maximum diameter ca 0-30-0-48. Promesonotal suture deeply impressed, metanotal 
groove varying from a feebly marked line to absent. In some specimens it is more distinct 
in the middle of the dorsum than laterally but in profile the metanotal groove makes at the most 
a very feeble impression in the dorsal outline and is often indiscernible. Propodeal dorsum 
with a narrow, median longitudinal groove, running from the metanotal groove to the declivity. 
A short, median, longitudinal groove is also present upon the posterior half of the pronotum, 
but this may be difficult to see in more heavily sculptured individuals. Node of petiole 
distinctly broader than long in dorsal view. Erect hairs variably present on dorsum of head 
and body, but apparently always absent from the propodeal dorsum and in many specimens 
also absent from the mesonotum. Full adult colour black. 

Sculpture very variable but mandibles, legs, antennal scapes and expanded lobes of frontal 
carinae always striate. Head and dorsum and sides of alitrunk always foveolate with striate 
interspaces, but the density of foveolae and intensity of striation variable. First and second 
gastral tergites foveolate, the foveolae varying in size, number and depth between individuals 
and the spaces between them usually feebly striate, although the intensity of striation varies 
from almost absent to very sharp. On the second tergite the anterior and posterior margins of 
the individual foveolae tend to be lost, and aligned foveolae run together to form a number of 
longitudinal impressions separated by raised welts, but again the formation of such sculpturation 
varies from individual to individual. 

Female. Answering to the description of the worker but with ocelli, and the alitrunk with 
flight sclerites. The queens tend to be somewhat larger than the workers of a given series, 
but their indices (CI and SI) fall within the worker range. 

Male. I have examined four male specimens which I tentatively associate with procerus 
as I have not been able to see the syntype males. These males differ markedly from the male 
associated with reichenspergeri in the shape of the subgenital plate. In reichenspergeri this is a 
narrow, broadly Y-shaped sclerite whereas in procerus it is short and broad, with short, thick, 
feebly projecting free corners which are slightly curved. The plate is thickly T-shaped, with 
the stem broad and the arms short. 

P. procerus is the most widely distributed species of the genus and, outside West 
Africa, is the species most commonly collected (foveolatus appears to be the com- 
monest species in West Africa) . P. procerus occurs throughout the forest zones of 
West and Central Africa and is also present in Uganda. Nest sites, as in foveolatus, 
appear to be situated in rotten wood. 



REVISION OF GENUS PS A LI DOMY RM EX 13 

I am not entirely convinced that the concept of procerus expressed above repre- 
sents a single species. The great variation seen in the workers could possibly 
conceal distinct but at present unrecognizable species as not enough material is 
presently available to show up any consistently differing characters which may be 
present. 

Material examined. 

Ghana: Tafo (B. Bolton); Tafo (D. Lesion); Kade (D. Lesion); Mt Atewa (D. 
Leston); Kukurantumi (D. Lesion). Uganda: Kawanda (M. M. Musoke & W. B. 
Banage); Kampala (C. C. Gowdey); Entebbe (C. C. Gowdey). Cameroun: Yole 
(G. Terron); Mt Nkolodon (G. Terron); no. 1070 and 1277 (no loc.) (G. Terron); 
Matute (B. Malkin). Gabon: Plateau dTpassa (/. A. Barra). Zaire: Leopoldville 
(Mouchet); Ituri Forest (C. P. Haskins). 



Psalidomyrmex wheeleri Santschi 

Psalidomyrmex wheeleri Santschi, 1923 : 263. Syntype workers, Zaire: Medje, Akenge and 
Niapu {Lang &- Chapin) (AMNH, New York; MCZ, Cambridge; MR AC, Tervuren; NM, 
Bulawayo) [examined]. 

Diagnosis of worker. Mandibles subtriangular. Expanded lobes of frontal carinae 
striate. SI 90 or more, DPI < 100. 

Further description. Worker. TL 13-6-14-6, HL 2-48-2-92, HW 2-20—2-60, CI 84-89, 
SL 2-00-2-52, SI 90-102, PW 1-80-1-96, PL 1-32-1-40, DPW 1-28-1-36, DPI 94-97 (8 measured). 

Answering to the description of procerus but a more slenderly built species 
with a narrower head, longer antennal scapes and a longer, narrower petiole. Maxi- 
mum diameter of eye in range 0-36-0-50. In general characters of colour, sculpture 
etc., wheeleri does not separate from procerus but the index ranges given below 
consistently differentiate the two species. 

wheeleri procerus 

Head narrower, CL range 94-89. Head broader, CL range 91-96. 

Scapes longer, SL range 90-102. Scapes shorter, SI range 80-88. 

Petiole longer than broad, DPI range Petiole broader than long, DPI range 

94-97. 106-118. 

The specimens constituting the type-series of wheeleri were first mentioned by 
Wheeler (1922) who identified them as procerus. This series consisted of nine 
workers from Medje, Akenge and Niapu, in Zaire, and were collected by the Lang & 
Chapin expedition. Wheeler noted that the specimens were all 'taken from the 
stomachs of toads (Bufo super ciliaris, funereus and polycercus) .' 

Santschi (1923 : 263) correctly decided that these specimens were not procerus 
but represented a new species, which he called wheeleri. 



i 4 B. BOLTON 

All the nine specimens mentioned by Wheeler were thus to be considered syntypes 
but to the present have not been labelled as such. I have traced eight of the 
nine and they are deposited as follows. 

AMNH, New York: 2 workers, one 'Niapu, Congo (H. 0. Lang) stomach Bufo 

polycercus f the other 'Medje, Congo (H. 0. Lang) stomach Bufo funereus.' (the 

latter now in BMNH). 
MCZ, Cambridge: 2 workers, one 'Medje, Congo (H. 0. Lang) stomach Bufo super- 

ciliaris'; the other 'Akenge, Congo [Lang) stomach Bufo polycercus.'' 
MRAC, Tervuren: i worker 'Musee du Congo; Ituri, Medje 1910 Exp. Lang, Chapin, 

stomach Bufo funereus.' 
NM, Bulawayo: 1 worker 'Akenge, Congo (Lang) stomach Bufo polycercus.'' 

I have added a circular, blue BMNH 'syntype' label to each of these specimens. 
The ninth and final syntype has not been located. 

A female in MCZ, Cambridge collection has been tentatively associated with these 
workers as it possesses the indices CI 89, SI 97, DPI 92, and otherwise generally 
resembles the workers. 

Material examined. 

Cameroun: Avom, 49 (G. Terron). Zaire: Banziville (Augustin); Ituri Forest, 
vie. Epulu (T. Gregg). 



acknowledgements 

I would like to express my thanks to the following, who have been of great help 
in sending me types and other material during the course of the study: 

Dr J. Decelle (MRAC, Tervuren); Mrs M. Favreau (AMNH, New York); Professor 
E. MeUini (IE, Bologna); Mr F. C. de Moor (NM, Bulawayo); Dr C. Baroni Urbani 
(NM, Basle) ; Mme J. C. Weulersse (MNHN, Paris) ; Ms J. C. White (MCZ, Cambridge). 



references 

Andre, E. 1890. Materiaux pour servir a la faune myrmecologique de Sierra Leone. 

Revue Ent. 9 : 311-327. 
Arnold, G. 1954. New Formicidae from Kenya and Uganda. Annls Mus. r. Congo Beige 

(N.S. in 4 ) Zool. 1 : 291-295, 6 figs. 
Bernard, F. 1952. La reserve naturelle integrate du Mt. Nimba 1. XL Hymenopteres, 

Formicidae. Mem. Inst. fr. Afr. noire 19 : 165-270, 15 figs. 
Bolton, B. 1974. A revision of the ponerine ant genus Plectroctena F. Smith. Bull. Br. 

Mus. nat. Hist. (Ent.) 30 : 309-337, 10 figs. 
Emery, C. 1901. Notes pour les sous-families des Dorylines et Ponerines (famille des 

Formicides). Annls Soc. ent. Belg. 45 : 32-54. 
Menozzi, C. 1922. Miscellanea mirmecologica. Annali Mus. civ. Stor. nat. Giacomo Doria 

(3) 9 [49] : 347-358, 4 figs- 



REVISION OF GENUS PS A LI DOM YR M EX 



15 



Santschi, F. 1913. Glanures de fourmis africaines. Annls Soc. ent. Belg. 57 : 302-314, 

5 figs- 

1920. Formicides nouveaux du Gabon, du Congo, de la Rhodesia et du Natal. Annls 

Soc. ent. Belg. 60 : 6-17, 3 figs. 
1923. Descriptions de nouveaux Formicides ethiopiens et notes diverses, 1. Revue 

Zool. Bot. afy. 11 : 259-295, 5 figs. 
1937- Fourmis du Congo et regions limitrophes. Revue Zool. Bot. afr. 30 : 71-85, 

6 figs. 

Wheeler, W. M. 1922. Ants of the Belgian Congo, part 2. Ants collected by the American 
Museum Congo Expedition. Bull. Am. Mus. nat. Hist. 45 : 39-269, 23 pis, 76 figs. 






Figs 1-6. 1, 2. Left mandible of (1) Psalidomyrmex reichenspergeri, (2) P. procerus. 
3, 4. Dorsal view of petiole and first gastral tergite of (3) P. reichenspergeri, (4) P. 
foveolatus. 5, 6. Outline of head of (5) P. sallyae, (6) P.feae ; antennae and pubescence 
omitted. 



l6 B. BOLTON 

INDEX 

Synonyms are in italics. 

clavicornis, G mabirensis, 8 

collarti, 12 obesus I2 

feae, 11 procerus, 12 

foveolatus, 7 . , 

reichenspergen, 

impressa, 1 1 sallyae, 9 

longiscapus, 12 wheeled, 13 



B. Bolton, B.Sc, A.R.C.S. 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 



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February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae. 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text-figures. December, 1973. £9-55- 

printed by Unwin Brothers Limited the gresham press old woking surrey England 



J GENERAL "* 
( f8APR!973 

THE BUTTERFLIES NAMED BY 

J. F. GMELIN 
(LEPIDOPTERA : RHOPALOCERA) 



R. I. VANE-WRIGHT 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 2 

LONDON : 1975 



THE BUTTERFLIES NAMED BY T. F. GMELIN 
(LEPIDOPTERA : RHOPALOCERA) 



*A*y 



BY 

RICHARD IRWIN VANE-WRIGHT 



Pp. 17-64 ; 6 Plates 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 2 

LONDON : 1975 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 2 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent. 

ISSN 0007-6431 



Trustees of the British Museum (Natural History), 1975 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 11 April, 1975 Price £yyo 



THE BUTTERFLIES NAMED BY J. F. GMELIN 
(LEPIDOPTERA : RHOPALOCERA) 



By R. I. VANE-WRIGHT 



CONTENTS 



Synopsis ........ 

Introduction ......... 

Historical review ........ 

Gmelin's method of numbering ...... 

a note on single type-specimens and lectotype designations 
The butterflies named by Gmelin from Museum Leskeanum 
Replacement names proposed by Gmelin .... 

Butterflies described in Museum Leskeanum but not named by 

Gmelin as new species 
Zschach, Cramer, Ambon and Surinam 
Summary of type-information 
Acknowledgements 
References ..... 

Index ..... 



Page 
19 
19 
20 

25 
26 
28 
53 

54 
55 
56 
57 
57 
63 



SYNOPSIS 

An historical review is given concerning the 26 butterfly names introduced by J. F. Gmelin. 
All these names appeared in the 13th edition of Linnaeus' Systema Naturae (1790), the great 
majority being based on previously unnamed descriptions of butterflies published by I. I. 
Zschach in Museum Leskeanum (1788-9). The fate of the Leske specimens is traced to the 
present day, and full synonymies are presented for all the species concerned. Two of the 
names are re-instated (at the subspecies level) giving a total of nine Gmelin names currently 
in use for recognized butterfly species or subspecies. Eleven new synonymies are established, 
four previous synonymies re-established, and 37 lectotypes designated (14 for Gmelin names, 
the remainder for taxa erected by other authors). In addition, neotypes are designated for 
three of the Gmelin species. Brief observations are given concerning the importance of Ambon 
and Surinam as type-localities, and the apparent survival of Cramer material in the British 
Museum (Natural History). 



INTRODUCTION 

This work deals primarily with the classification of the 23 butterfly species named 
in 1790, by J. F. Gmelin, from descriptions in the catalogue of the N. G. Leske 
collection (Karsten, 1789; see PI. 1, fig. 2). Gmelin published these names in the 
13th edition of Linnaeus' Systema Naturae, of which work Gmelin was effectively 
both author and compiler. The 13th Systema Naturae includes a large number 
of new names, normally attributed to Gmelin in Linnaeus, the great majority of 
which were coined by Gmelin for previously unnamed animals, plants and minerals 



20 R. I. VANE-WRIGHT 

already described or illustrated in the literature. Sherborn (1902) lists all the 
Gmelin animal names, but in many groups of insects they have been almost com- 
pletely ignored (Dipt era), or partially ignored (Coleoptera, Orthoptera). The 
butterflies, however, have been largely accounted for, at least from a nomenclatorial 
standpoint, due to a series of chance events. Almost all were named from the 
Leske collection, which was shortly afterwards purchased by the then Dublin 
Society. Over 50 years later the remains of the collection came to the attention 
of W. F. Kirby, who was then working in Dublin. Kirby (1869; 1871; 1877), 
who at that time was preparing his pioneer butterfly catalogue, drew attention 
to the general problem of the Gmelin names, at the same time accounting in detail 
for the majority of the butterfly species involved. As already indicated, however, 
in many groups of insects the Gmelin names are still ignored, even for the purposes 
of primary homonymy. Since the classification and nomenclature of butterfly 
species has reached a relatively advanced stage, it seems appropriate, some 100 
years after the appearance of Kirby's catalogue, to review the Gmelin butterfly 
species again. In doing so, some fresh insight has been gained concerning the 
problems of the Gmelin names and the Leske collection. It is hoped that the present 
paper will be of use not only to butterfly workers, but also to those who may wish 
to tackle the problems raised by Gmelin names in other groups. 



HISTORICAL REVIEW 

In 1788 G. Muller of Leipzig published a work by I. I. Zschach, the Pars Ento- 
mologica of Museum Leskeanum (PI. 1, fig. 1). The latter (PI. 1, fig. 2) was to be 
a catalogue of the collection of natural history objects, animal, plant and mineral, 
formed by Nathaniel Gottfried Leske. Of Russian parentage, Leske was living 
and working in Leipzig at the time of his death in 1786. He described himself 
as a 'natural historian and economist', being the author of a considerable number 
of popular articles, books and translations. 

The first complete part of Museum Leskeanum, the Regnum Animate (PI. 1, 
fig. 2), was published by Muller in the following year, 1789. This work, largely 
written and edited by D. L. G. Karsten of Marburg, included Zschach's Pars Ento- 
mologica as 'Classis V. Insecta' (PI. 1, fig. 3), which comprised pages 1-136, the 
original pagination of Zschach. Classes I-IV take up pages I-XLIV, following 
Karsten's preface, etc.; the pagination of the work is thus rather complex, due to 
the prior publication of Zschach's contribution, the same type settings being used 
for both. It is interesting to note that Class VI, Vermes, follows on from the Insecta, 
as page 137, etc. 

Karsten's contributions to Museum Leskeanum are based on the Linnaean system 
of classification, but for the insects Zschach followed Fabricius (PI. i, figs 1, 3). 
Zschach evidently considered the Leske collection to contain a large number of 
new species, or, perhaps more accurately, species which he could not recognise 
amongst the descriptive literature known to him. Whilst the known species were 
merely listed by Zschach (e.g. PI. 2, fig. 5, species 1-5), these unknown species 
were described, but names were not given to them (e.g. PI. 2, fig. 5, species 6; 






BUTTERFLIES NAMED BY GMELIN 21 

PI. 3, fig. 6). Thus in the Pars Entomologica each previously named species of an 
order is given a consecutive number (starting from i), then an abbreviation indicat- 
ing the section of the genus, then the specific epithet, and finally a literature reference. 
The unnamed species are numbered, but appear without a binomen or reference. 
Instead, after the abbreviated indication of the generic section, an asterisk appears, 
followed by Zschach's Latin description of the insect concerned (PL 2, fig. 5; 
PI. 3, fig. 6). This pattern of presentation is followed for the hundreds of unnamed 
insects thus described by Zschach in Museum Leskeanum, all of which may be 
readily recognized by the asterisk mark. 

In 1790 (Hopkinson, 1908), volume 5 of the 13th Linnaean Systema Naturae 
appeared, compiled by J. F. Gmelin, and also published in Leipzig. Within two 
to three years Gmelin's edition was entirely reprinted verbatim by Delamolliere 
at Lugduni, i.e. Lugduni Monachorum (not Batavorum), or Lyon (not Leiden). 
The format and pagination were identical with the original, but the type was reset 
and, although very accurate, the occasional misprint occurs. Since volume 1 
(Animalia) had only one title-page for its seven bulky parts, only part 1 is identifiable 
at sight, each other part having simply its own half-title. Part 5, Insecta (1792) 
can be identified only by the misprints. So far only three of these have been 
noticed, all affecting the numbering of Papilio species: p. 2248 numbers run 878, 
351, '44', 44 (they should be . . . 45, 44); p. 2250, number '336' should be 356; p. 2360, 
number '701' should be 901. Otherwise either printing appears satisfactory for 
study, provided one knows which is used. Since the bibliographic data for the 
reprint are identical, apart from the date and publisher's detail, with Gmelin's 
original, only the latter is listed in the references. 

In the 13th Systema Naturae Gmelin listed all the published names known to 
him. In addition he named many new species, almost invariably it seems by 
reference to already published descriptions or illustrations of unnamed organisms, 
bestowing names on these for the first time. Gmelin also introduced a number 
of replacement names, where homonymies came to his attention (e.g. Papilio 
philetes Gmelin, 1790 : 2364). 

One such work to which Gmelin consistently referred was Museum Leskeanum, 
and he thus gave names to the great majority (but not all) of the unnamed insects 
described by Zschach. For each new species so named, Gmelin gave a page refer- 
ence to the description in Museum Leskeanum, the Zschach species number, and 
a virtual copy of the latter's original description. To illustrate this we may take 
Zschach's Papilio 46c, which appears on page 89 of Museum Leskeanum, as the 
fourth species of section 6, the Danai festivi (PL 3, fig. 6). This should be compared 
with the entry for Papilio no. 887 (PL 3, fig. 7) which appears on page 2289 of 
Gmelin in Linnaeus, under the Danai festivi. The latter is the original description 
of Papilio claviger Gmelin; it is clearly drawn directly from Zschach's Papilio 46c, 
Gmelin's description being a barely altered paraphrase. 

In 1792 the Dublin Society purchased the Leskean mineral cabinet (for a sum 
of about £1350; Berry, 1915 : 156); evidently at the same time they also received 
a large part, if not all, of the Leske zoological and botanical collections (White, 
191 1 : 8). It thus seems possible that a prime purpose of Karsten's work in cata- 



22 R. I. VANE-WRIGHT 

loguing the Leske collection was to facilitate the sale of the material, at the same 
time providing a valuable record of a notable museum collection. According 
to Berry (loc. cit.) the Leske mineral collection was considered one of the finest 
in Europe in its day, Karsten also being an outstanding mineralogist. Berry 
unfortunately gives scant information on the Leske zoological material. 

In 1813 a catalogue of the natural history specimens in the Dublin Society museum 
was published; the title page is shown in PI. 1, fig. 4. It is evident from the short 
'Advertisement' by Bernard O'Reilly which prefaces the work (PI. 4, fig. 10), and 
also from some of the finer points in the work itself, that whoever made the catalogue 
was well versed in the systematic natural history of the time. Reference to the 
works of White (1911), Berry (1915), Praeger (1949) and a number of manuscripts 
and publications available in the National Museum of Ireland, has shed no light 
on the catalogue, or on Bernard O'Reilly. Unless proven otherwise, it seems 
practical to attribute this obscure work to the equally obscure O'Reilly. White 
(191 1 : 10) even goes so far as to say that a directive issued in 1826 to Giesecke 
to collect insects in county Donegal is 'the first mention of an Entomological Collec- 
tion in the Museum', when O'Reilly in fact, 13 years earlier, had already listed over 
2500 species of insects in the Society's collections! Later, however, White (op. cit.: 
14) probably unwittingly refers to O'Reilly's catalogue ('In 1843 the society called 
for a report ... it refers . . . also to a Zoological Collection (Leskean) and the cata- 
logue thereof). The only clue to O'Reilly is a book published in 1818 {Greenland, 
the adjacent seas, . . .) by someone of the same name; PI. 4, fig. 12 shows the entry 
for this work in the British Museum (Natural History) catalogue (1910 : 1476), 
and the vitriolic comment reprinted from the London Quarterly Review which 
accompanies it. Possibly the publication of this book so embarrassed the Dublin 
Society that all account of O'Reilly was banished from their records. Evidence 
that the two books are by one and the same O'Reilly is to be found on pages 97-98 
of the 'Greenland' volume, where it is stated that 'the arrangement of such animals 
as I have seen is conformable to the system of Linnaeus, according to the last 
edition of his celebrated work by Gmelin'. In any event, it is highly unusual to 
find anyone with a clearly Irish name being credited with any contribution towards 
the Society's works at that time, the organization then being firmly in the control 
of the English and Anglo-Irish. 

However, comparison of O'Reilly's catalogue with Museum Leskeanum indi- 
cates that the Dublin Society insect collection in 1813 was very largely made 
from the Leske collection, which they had purchased 21 years previously. The 
species numbers and names often correspond exactly with Zschach's work, with 
the interesting addition of Gmelin's names where Zschach had deferred to name 
them. At least on some occasions where Gmelin had ignored certain Zschach 
descriptions, the insect concerned is referred to as a variety of one of the others 
listed. In the great majority of cases, where in Museum Leskeanum the locality 
is given as 'Exoticus', in O'Reilly's catalogue this is rendered (needless to say, usually 
incorrectly) as 'India'. We may compare the entry shown on page 75 of O'Reilly 
for Papilio 46c (PI. 3, fig. 8) with the corresponding entries in Zschach and Gmelin 
(PI. 3, figs 6, 7). Thus O'Reilly gives for each such species the Museum Leskeanum 



BUTTERFLIES NAMED BY GMELIN 23 

number of Zschach (which, according to the preface, would have been 'affixed to 
the subject' -see PI. 4, fig. 10; this is largely confirmed by Kirby, 1869 : 355), 
the Gmelin name, an English name, and a rendering of 'exoticus' as 'India 1 . This 
pattern is repeated throughout, with a few minor changes. Perhaps the most 
interesting variation in the butterflies concerns the first species named from the 
Museum Leskeanum material - no. 6 (PI. 2, fig. 5). Gmelin called this Papilio 
argyrios, but O'Reilly lists this as '6. P. E. A. Leskii. Leskean Butterfly. India' 
(PI. 4, fig. 11), this being one of a few species in the catalogue so named after Leske, 
and which seem to constitute the only recognition of the original source of the 
bulk of the collection. However, O'Reilly's catalogue appears to provide conclu- 
sive proof that the types of the great majority of those Gmelin species, based on 
the descriptions of insects by Zschach in Museum Leskeanum, reached Dublin 
and were extant in 1913. 

W. F. Kirby, who joined the staff of the, by then, Royal Dublin Society in 1867 
(Praeger, 1949 : 116; not 1865 as suggested by White, 1911 : 25), presented a 
paper in 1869 which dealt exclusively with the forgotten works of Gmelin and 
Zschach (strangely, he also makes no mention of O'Reilly), with the express purpose 
of generally drawing attention to them, and in particular to provide information 
about the butterflies (e.g. PI. 3, fig. 9 shows Kirby's entry for Papilio 46c of Zschach). 
Kirby was an Assistant-naturalist for 12 years at Dublin, and in this paper he notes 
'the remains of Leske's collection are in the Museum of the Royal Dublin Society, 
but the greater number of specimens have succumbed to the ravages of time and 
neglect'. Kirby lists 25 (correctly 24) butterflies as named by Gmelin, 23 of which 
were drawn from the 25 species described by Zschach (Kirby wrongly attributes 
Papilio sectator Meerburgh to Gmelin). At this time Kirby had evidently located 
type-material of some 16 of the latter. At the end of the paper he makes the 
following comment of general interest: Tmayadd that Leske'scollection was purchased 
for the Royal Dublin Society at the end of last century. The Lepidoptera Heterocera 
are all destroyed or seem unrecognisable. There are, however, a good many Coleop- 
tera, &c, still in existence.' 

While still at Dublin, Kirby produced his Synonymic Catalogue of Diurnal Lepi- 
doptera (1871), and also his Supplement to this work (1877). Embodied in these 
are the results of his 1869 paper on the Gmelin names, and also a number of further 
discoveries or conclusions about this problem which he must have made between 
1869 and 1877 (e.g. PI. 5, figs 13-16, shows various entries concerning Papilio 
46c). In no case is any further explanation given; as throughout the catalogue, 
the 'facts' are indicated by bold statement, the surmises or uncertainties by '?'. 
Shortly after leaving Dublin for London, Kirby (1880) published a catalogue of 
part of the Lepidoptera collections, as he had left them, at Dublin; strangely, 
despite the seemingly full treatment given to the butterflies, no mention is made 
of any Leske specimens or the Gmelin types. The now National Museum of Ireland 
possesses an interleaved copy of Zschach (1788), annotated in Kirby's handwriting 
(PI. 5, figs 17, 18); as surmised above, Kirby evidently did do further work on 
the Leske material after 1869, the results being included in his catalogue, or its 
supplement, at various points. 



24 R. I. VANE-WRIGHT 

Despite Kirby's hope of stimulating interest in Gmelin, this difficult area of 
the early literature has still received scant attention from entomologists, probably 
understandably. The great majority of Gmelin insect species must remain 
unidentifiable, except in the few instances where type-material may still exist (as 
it does with at least some of the species named from the Leske material) . Probably 
the single major taxonomic problem, however, concerns primary homonymy; 
all the Gmelin names appear to have been noted by Sherborn (1902), in his Index 
Animalium, but other cataloguers have taken varying account of them, some 
ignoring them, others including them. Most curiously, Kirby himself, in his 
Orthoptera catalogues (published long after his days in Dublin), includes some but 
not all of the Gmelin species. 

Those studying the butterflies have been relatively fortunate in this regard, 
due to the high proportion of types remaining, and the excellent work of Kirby. 
Even here, however, little cognisance of Kirby's work and how he was able to achieve 
his results seems to have been made by most workers, except occasional British 
entomologists. As an example of the former we may quote Weymer (1910 : 182, 
footnote), writing in 'Seitz': 

'Whether Antirrhea bifasciatus Gmelin cited by Kirby in his Catalogue really 
belongs to the genus Antirrhea or even to the American fauna, appears question- 
able, as it cannot be recognised from the description and the locality was not 
known to the author. The species is only designated as exotic, and hence may 
just as well belong to the Indian or African fauna. The original of the description 
was in the Museum Leskeanum, and no further example is known. According 
to this description . . . Habitat extra-European (Gmelin, Syst. Nat.)'. 

Concerning bifasciatus, Kirby (1869 : 360) writes: 'One of the Satyrinae. A 
fragment of one of the types is still in existence; but I have not yet succeeded in 
identifying it.' Later (1871 : 642), in the Appendix to his catalogue, Kirby indi- 
cates that it is a species of Antirrhea, placing it after A. taygetina Butler. In the 
Dublin interleaved copy of Zschach (1788), opposite page 91, the following note 
in Kirby's handwriting appears: '59 P.P. Bifasciatus, Gmel. Syst. Nat. I. 5 
p. 2290 n. 893. = Antirrhaea sp.' (PI. 5, fig. 17). 

Sadly, the 'fragment' of bifasciatus is no longer to be found in Dublin. However, 
a number of factors support the possibility that Kirby was right in his placement 
of this species. First, as will be evident from the list below, Kirby was essentially 
correct in all his firm identifications of Leskean/Gmelin types, where this has been 
confirmed by their continued existence. Secondly, where there was doubt about 
either the types or the placement of the species he always indicated it. Thirdly, 
the genus Antirrhea is a very characteristic S. American group; Kirby (1880 : 295) 
indicates that the Dublin Museum then possessed four species of Antirrhea, all 
of which are still present and were clearly correctly identified by him. Fourthly, 
Kirby did not just add the species to the genus, but placed it to follow a well defined 
species, suggesting that possibly he had been able to do more than just recognize 
it as a member of Antirrhea. Against this it is fair to say that the descriptions of 
Zschach and Gmelin do not fit any species of Antirrhea currently recognized; how- 



BUTTERFLIES NAMED BY GMELIN 25 

ever, the majority of Antirrhea species are rare in collections, and at least three 
are still only known from uniques, quite apart from bifasciatus (Vane-Wright, 
unpublished). Thus while it is frustrating that the fragmented specimen which 
convinced Kirby to place bifasciatus in Antirrhea no longer exists, it would seem 
that Weymer did not have all the facts at his disposal, and that it may be more 
prudent to follow the excellent Kirby in this regard, leaving bifasciatus Gmelin 
as a species inquirenda of the genus Antirrhea. 

As a result of examining the collections at the National Museum, Dublin, I 
am able to present a revised version of Kirby's 1869 paper, including a large number 
of further references, lectotype designations, and modern indications of classifica- 
tion. While this will not form the last word on the subject, I hope that it will 
settle the majority of outstanding problems concerning the Gmelin butterfly names, 
bringing together most of the relevant information, while also stimulating others 
to tackle at least the problems of primary homonymy caused by the Gmelin names 
in their respective groups. 



GMELIN'S METHOD OF NUMBERING 

Gmelin's numbering of Papilio species is bewildering at first sight, but actually 
is logical and rather neat. His system, which probably applies to all large genera 
and was doubtless evident to his contemporaries, is worth examining. 

His first six species (pp. 2225-2226) are nos. 2, 274, 3, 275, 276, and 4. On page 
2248 the numbers run 878, 351, 45, 44, and his final page 2370 carries nos. 875, 876, 
268-271, 877, 272, 273. One number is misprinted: '550' for 558 (p. 2301). This 
reduces to three series of intercalated numbers: 1-273 (omitting 112, P. enceladus, 
an oversight?); 274-877 (omitting 345, 622); and 878-901 (omitting 884). The 
last series runs serially, but the first two are only approximately so. The first 
series starts with nos. 2-10, 1, 11-19, 25 . . . and ends with nos. 256-273 (no. 255 
on p. 2287 lying between nos. 128-130, 239, 240, 501, 502, 242, 503, 504 and nos. 
505-511, 262, 512-515, 885-893, 131). 

Gmelin was revising Linnaeus (1767), where the genus Papilio included 273 
species. To refer back, Gmelin simply used the Linnaean numbers, so his species 
1-273 immediately equate with those of Linnaeus, and all except one bear the 
same names. Their sequence is varied to accord with his new classification require- 
ments, and additions (the second and third series) are interpolated in their proper 
places. This economical method explains why Gmelin gave no other clarification 
when he substituted for no. 235 P. pirithous Linnaeus, 1767, the replacement name 
P. barbarus, no. 235, on page 2352. No reason for replacing the perfectly valid 
Linnaean name is apparent, but it was probably in deference to Fabricius, whose 
junior homonym P. pirithous Fabricius, 1775, for an entirely different species he 
had already cited on page 2281, no. 478. In any case P. barbarus Gmelin is clearly 
an invalid junior objective synonym of P. pirithous Linnaeus, 1767. 

Apart from that exception in Gmelin's first series there are only a few minor 
name changes. An interesting case is P. ancaeus Linnaeus, 1758 (and 1767, no. 
184), which Gmelin (p. 2276, no. 184) notes had been misspelt ancaea by Cramer. 



26 R. I. VANE-WRIGHT 

To make this abundantly clear he gives the Linnaean reference and number, which 
is also helpful because the preceding number is 113 and no. 114 is not found until 
the next opening, while nos. 183 and 185 do not appear until pp. 2320 and 2321. 
However, as will be seen later, the presence of no. 184 at this point probably caused 
a misnumbering in the final sequence. 

Gmelin's second sequence, nos. 274-877, comprises all other names published 
to that date. All but two of these had been included in the latest definitive revision 
of the Insecta, by Fabricius (1787), and Gmelin gives references to that work for 
each species. Fabricius listed 834 species, but overlooked or merged some 38 
of those recognized by Gmelin and Linnaeus, which accounts for most of the dis- 
crepancy between 834 and 877. The balance is due to the omission by Gmelin 
of three numbers (112, 345 and 622), and his inclusion of two further species: 
n0 - 355. P- sectator Meerburgh, 1775, and no. 601, P. ceres Fabricius, 1775. 

The final 23 species, which comprise the series nos. 878-901 (omitting 884), 
were newly named by Gmelin, and are discussed below. The reason for the omission 
of no. 884 was perhaps that no. 883 came early in the Danai candidi on p. 2261, 
while the next group of the series was not reached until the end of the Danai festivi 
on p. 2289. While looking back to establish the number at the latter point, the 
number out of sequence to catch the eye was, as noted above, no. 184. Gmelin 
may have misread this as 884 and then proceeded with no. 885. 

A NOTE ON SINGLE TYPE-SPECIMENS AND LECTOTYPE DESIGNATIONS 

An ambiguity has arisen from certain wording in the International Code of Zoological 
Nomenclature, regarding the interpretation of holotypes and syntypes, and the 
necessity or otherwise of fixing lectotypes in certain instances. This concerns 
the situation where a species is described from an unstated number of specimens, 
with no original published designation of a specific 'type' or holotype, and where 
now only one original specimen is to be found. Some workers, perhaps a majority, 
regard such specimens as 'unique' types, and so treat them as holotype specimens. 
Dr Roger Crosskey has put forward an eloquent case in the hope of formalising this 
'holotypist' approach; the reader is referred to the statement by Crosskey (1974 : 272) 
for details of the situation and his argument. However, the present author has 
always used the alternative interpretation: if there is no objective evidence in an 
original description as to whether the original type-series was multiple or not, then 
in such cases, even if only one original specimen can now be found, it requires 
eventual fixation as a lectotype. In the meantime such a specimen remains a 
type of indefinite status, or may be regarded as a lone syntype in a series of unknown 
size (which includes, of course, the implicit possibility that the original series did 
only consist of one specimen). 

My interpretation of the Code is based on the operational principle that if it can 
be unequivocally inferred from the original description that there is a unique or 
'type' specimen, then that specimen is the holotype, but in all other cases, to provide 
a specified unique type for the taxon concerned, a subsequent lectotype designation 
is necessary, even if only a single original specimen can now be found. 



BUTTERFLIES NAMED BY GMELIN 27 

Crosskey (op. cit.) lists a number of objections to such a procedure, or at least 
to the designation of surviving unique specimens of the nature under discussion, 
as lectotypes. One objection is that other workers, seeing a lectotype to have been 
fixed, will expect at least one paralectotype to exist also. My use of the term 
lectotype is to indicate a clear fixation of a unique type-specimen to serve for 
nomenclatorial purposes, where no such fixation is explicit or implicit in the text 
of the original description concerned. In designating unique specimens of the sort 
under discussion as lectotypes, it should always be indicated that only the single 
specimen is known to exist, and that the size of the original series was quite 
unspecified. 

Another point advanced by Crosskey is that, especially with older authors where 
most of these problems occur, it is rare to find more than one surviving type-specimen, 
suggesting that most old taxa were based on single specimens. This may be very 
true in a group like the Diptera, but it is a much less safe assumption with the 
Lepidoptera. Most of the taxa dealt with in this paper are of considerable antiquity. 
Analysis of the original descriptions of 56 species noted here reveals the following: 
only three have explicit holotypes; 38 are based on a completely undefined number 
of specimens; n are clearly based on series of more than one specimen, but the 
number of specimens is in no way indicated; and four are based on syntypic series 
with the included number of specimens being indicated. Of the 38 taxa with an 
undefined number of specimens in the original series, I have been unable to trace 
any type-specimens in 34-2 per cent, of cases, one specimen in 55-3 per cent, of 
cases, and two or more type-specimens in 10-5 per cent, of cases. Of the 11 species 
with unstated, but clearly multiple original series, I have been unable to find any 
type-specimens in 45-5 per cent, of cases, one specimen in 27-25 per cent, of cases, 
and more than one in 27-25 per cent, of cases. I think these figures show that 
in the case of butterflies, a large number of errors would be made if it were regularly 
assumed that a single surviving specimen indicated that the taxon in question 
was originally based on that specimen alone. With butterflies, the long history 
of private collectors, private loans, purchases, auctions, removal of labels, and even 
theft, has conspired to cloud the picture with uncertainty. In addition, some 
authors (e.g. Hans Fruhstorfer) are bemusing in their inconsistency with regard 
to their original publications, and the retrospective criteria now applied to decide 
type status. In this context, one need only think of the nightmare created by 
'Seitz' for the conscientious museum worker concerned about applying the correct 
status to the types in his care. 

In a final, and most significant argument, Crosskey claims that by designating 
a unique specimen as a lectotype, one is 'unnecessarily tying the hands of future 
zoologists', should other original specimens later be found to exist. Certainly, 
lectotype designations are final, if correctly made, but I believe that publishing 
accounts in which supposedly unique specimens are stated to be holotypes is opera- 
tionally equally final. Such 'holotype designation' would always tend to be regarded 
as a 'previously valid type restriction' by subsequent workers. However, the 
whole question of restricting the freedom of future zoologists raises a more funda- 
mental problem and division. The present author is a somewhat unwilling convert 



28 R. I. VANE-WRIGHT 

to the 'lectotype designation cult'. My original viewpoint was that if a syntypic 
series existed, and if in the opinion of a subsequent worker all the syntypes applied 
to the same taxonomic grouping concept, then a lectotype designation served no 
useful function, but was indeed undesirable, since it tied the hands of future zoologists. 
Should such a series at a later date be deemed to be mixed (rare for butterflies, 
common for Diptera), then the zoologist concerned would be free to make the most 
appropriate choice in selecting a lectotype. However, I am now largely persuaded 
that, provided lectotype designations are not made casually (as for instance in 
type-catalogues, or figure legends, both frequent and regrettable practices), it 
is better that a taxonomist in doing work of a revisional nature should unequivocally 
fix definite, single type-specimens for all taxa in question, wherever possible, so as 
to reduce present and future uncertainty to a minimum. I am prepared to accept 
this in the current type-oriented approach to zoological nomenclature, since it 
seems inconsistent to advocate all new taxa to have fixed holotype specimens, 
yet wish to retain old type-series wherever possible as syntypic, 'just in case'. In 
fact, if we are seriously to consider flexibility for future zoologists, it would arguably 
be better to abandon the holotype concept altogether, and introduce an extended 
lectotype concept for use where necessary (this could include the neotype concept) ; 
but the present system has become far too well entrenched for the present author 
to have any great desire to see it changed. 

Accepting the present system, and as an 'unwillingly converted lectotyper', 
I therefore regard the proposal advanced by Crosskey as neither fitting with one 
ideal nor the other. If lectotypes are to be designated as a matter of course where 
syntypic series exist, regardless of any doubt as to the conspecificity of the specimens, 
then in my opinion lectotypes should also be fixed for single extant specimens where 
there is no indication of unique status in the original description, provided this 
is done explicitly, and only in taxonomic treatments of an essentially revisional 
nature. 

In conclusion to this discussion, I think it fair to say that in practice the two 
interpretations will nearly always lead to the same result. Rather than press for 
uniformity on a matter like this, I believe that it may be better not to tie the hands 
of present zoologists, allowing them the flexibility to apply whichever procedure 
seems the most sensible to them. What seems sensible in one group, and not so 
sensible in another, will depend largely on the history of collecting and museum 
techniques previously applied, and also on the nature of the particular organisms 
in question. 

THE BUTTERFLIES NAMED BY GMELIN FROM MUSEUM LESKEANUM 

The Gmelin butterflies named by reference to Museum Leskeanum are dealt 
with in their original order of presentation. For each, the presently accepted name 
is given centrally, in bold type, together with an indication of its family, followed 
by full species and important literary synonymy. Under a heading giving the 
original Gmelin name, there follows a brief taxonomic discussion in each case, with 
observations on the most likely type-locality, and including either a lectotype 



BUTTERFLIES NAMED BY GMELIN 29 

or neotype designation where appropriate. In a number of instances lectotypes 
are also designated for synonymous taxa described by other authors. 

The abbreviations listed below are used for depositories of type-material. 

BMNH British Museum (Natural History), London, England. 

CM Carnegie Museum, Pittsburgh, U.S.A. 

LC Linnaean Collection, Linnean Society of London, England. 

MLU Museum Ludovica Ulrica, Uppsala, Sweden. 

MNHN Museum national d'Histoire naturelle, Paris, France. 

MNHU Museum fur Naturkunde der Humboldt-Universitat, Berlin, D.D.R. 

NMI National Museum of Ireland, Dublin, Eire. 

RNH Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands. 

RSM Royal Scottish Museum, Edinburgh, Scotland. 

USNM National Museum of Natural History, Washington D.C., U.S.A. 

Zl Zoological Institute, Leningrad, U.S.S.R. 

Alcides orontes (Linnaeus) [Heterocera: Geometroidea, UraniidaeJ 

Papilio orontes Linnaeus, 1763a : 19. [Ambon] (MLU). 

Papilio orontes Linnaeus, 17636 : 402. 'In India'. 

Papilio orontes; Clerck, 1764 : pi. 26, fig. 1. 

Papilio Equites Achivi sp., Zschach, 1788 : 87, no. 6*, pi. 2, fig. Glossata 6. 'Exoticus'. 

Papilio argyrios Gmelin, 1790 : 2248., no. 878. 'Extra Europam' [? Ambon] (formerly in 

NMI; type-material destroyed). Synonymy by Kirby, 1892 : 16. 
Papilio leskii O'Reilly, 1813 : 73, no. 6, nomen nudum. 'India'. [This name is objectively 

synonymous with argyrios Gmelin, being based on the same material.] 
Alcides orontiaria Hiibner, [1822] : pi. [218], figs 3, 4. [No locality.] (Type-material lost.) 

Synonymy from Dalla Torre, 1924 : 6; correct date: Hemming, 19376 : 149. 
Alcidis orontiaria Hiibner, [1823] : 289. Correct date: Hemming, 19376 : 149. 
Papilio argyrios Gmelin; Kirby, 1869 : 356. 
Nyctalemon argyrios (Gmelin); Kirby, 1871 : 638. 
Alcides orontes (Linnaeus); Kirby, 1880 : 335. 
Alcidis orontes (Clerck); Kirby, 1892 : 16. 
Papilio orontes Linnaeus; Sherborn, 1902 : 702. 
Alcidis orontes (Clerck); Dalla Torre, 1924 : 6; Seitz, 1929 : 94. 

Papilio argyrios Gmelin. I have been unable to trace any type-material in 
Dublin. Kirby (1880 : 335) lists one specimen of Alcides orontes Linnaeus, which 
is extant, but lacks data. It seems unlikely that this specimen was part of the 
Leske collection; curiously, no Leskean butterfly specimens are listed anywhere 
in Kirby's 1880 paper, and presumably the same would apply to this moth species. 
A good original figure is given by Zschach (1788 : pi. 2, fig. Glossata 6); the same 
plate is reproduced in Karsten (1789). The original most probably came from 
Ambon, and argyrios may thus be regarded as a strict synonym of orontes Linnaeus 
(Corbet, 1949 : 192-193, summarises information relevant to determining the 
type-locality of orontes). 

Papilio leskii O'Reilly, noted in the synonymy above, is best regarded as a 
nomen nudum. If, however, it is treated as an available name, it falls as an objec- 
tive synonym of argyrios. 

Some confusion appears to exist in the literature concerning the authorship, 
date and even genus of orontes Linnaeus. The question of the spelling Alcides 



3 o R. I. VANE-WRIGHT 

v. Alcidis appears to have been settled finally by Hemming (1937& : 149), in favour 
of the former. The authorship and date problem is more vexed. A number of 
workers have attributed the name orontes to Clerck, 1759. It would appear, how- 
ever, that these workers overlooked the division of Clerck's I cones into two parts; 
the title page of the second part, in which the original iUustration of orontes appears, 
is dated 1764. The earliest dated work to mention orontes is the dissertation 
delivered by Boas Johansson in 1763, Centuria Insectorum Rariorum (Linnaeus, 
1763a), which was also reprinted the same year in Amoenitates Academicae (Lin- 
naeus, 17636). It is considered normal to regard the authorship of new names 
in such dissertations as being that of Linnaeus, who was apparently responsible 
for most of the content of these works. Both these 1763 publications, however, 
make clear reference to 'Clerk t.26 f.l'; Higgins (1970) has suggested from other 
evidence that the component plates of Clerck's work may have been issued con- 
tinuously, perhaps separately, from 1759 to 1764. It might therefore be correctly 
interpreted that Clerck had already published orontes prior to Johansson and 
Linnaeus. It would seem preferable, however, in the absence of proof to the con- 
trary, to continue to regard the proper date of publication of the second part of 
Clerck's Icones as 1764; prior 'publication' probably consisted of private circulation 
of the separate plates, as they were completed, among friends and co-workers. 
It may also be noted that Hemming (1967 : 288, entry for Metamandia) was of 
the categorical opinion that all names published in Clerck's Icones should be 
attributed to Linnaeus. 

I have followed Seitz (1929) with regard to the determination of these moth species, 
but any future reviser should carefully consider the type-material and original 
figures of orontes, argyrios and orontiaria. 



Pierella hyalinus hyalinus (Gmelin) [Satyridae] 
(PI. 6, fig. 19) 

[Papilio Una Linnaeus; Cramer, [1780] : 5, pi. 291, figs A, B. 'Suriname'. Misidentification.] 

Papilio parnassi sp., Zschach, 1788 : 88, no. 26*. 'Exoticus'. 

Papilio hyalinus Gmelin, 1790 : 2259, no. 879. NEOTYPE $, Surinam (BMNH), here 

designated [examined]. 
Papilio hyalinus; O'Reilly, 1813 : 74, no. 26. 'India'. 
Papilio hyalinus Gmelin; Kirby, 1869 : 356; 1871 : 637. 
Pierella hyalinus (Gmelin); Kirby, 1877 : 698; Kirby, 1880 : 295. 
Pierella hyalinus hyalinus (Gmelin); Brown, 1948 : 63. 

Papilio hyalinus Gmelin. No original type-material seems to have survived 
at Dublin, evidently having been destroyed sometime after 1813, and before Kirby's 
arrival in 1867. In the interleaved copy of Zschach at Dublin, Kirby first entered 
'? = Antirrhaea philoctetes L.'; subsequently he deleted this and wrote ' = Pierella 
dracontis\ The most probable original type-locality for hyalinus is Surinam. 

Kirby (1877 : 698) placed dracontis Hubner ([1819] : 53) as a junior synonym 
of hyalinus. Subsequently Brown (1948) has used these names, respectively, 
for the Amazon and Guyana region subspecies of hyalinus which he recognized. 



BUTTERFLIES NAMED BY GMELIN 31 

Brown (op. cit.) gives good reasons for believing the type-locality of hyalinus to 
be Surinam. He also discusses the early confusion between lena Linnaeus and 
hyalinus, and the later application of the name hyalinus by other workers to insects 
from the Amazon and Trinidad. Brown indicates the illustration by Cramer 
(1780 : pi. 291, figs A, B) of a butterfly from Surinam (misidentified by Cramer 
as lena) to be the earliest description of this species as he interprets it; the specimen 
believed to have been illustrated by Cramer is now in the BMNH (see the general 
notes on Cramer, p. 55). Apart from the earlier confusion over species and sub- 
species names, Mr M. P. Clifton (unpublished work) is of the opinion that a complex 
of sibling species is involved in this group. In order to stabilize existing usage, 
the male specimen, believed to have been illustrated by Cramer as referred to above, 
is hereby designated neotype of Papilio hyalinus Gmelin. It bears the following 
labels: /Neotype/Surinam Coll. Lenep/Dracontis Hiib./H. Dracontis Hub Lena 
Cramer/Felder Colin. /Rothschild Bequest BM 1939-1/Papilio hyalinus Gmelin, 
det. R.I.V-W./, and is illustrated on PL 6, fig. 19. 

In this context it may be interesting to note Papilio sectator Meerburgh (1775 : 
pi. 10, [83]). Kirby (1869 : 356) attributed this Meerburgh name (but not others) 
to Gmelin (1790 : 2250), perhaps assuming that Meerburgh had illustrated the 
species but not named it. Kirby (1871 : 642) later corrected this mistake, noting 
sectator Meerburgh as a junior synonym of true Pierella lena (Linnaeus). 



Delias isse isse (Cramer) [Pieridae] 

Papilio isse Cramer, 1775 : 87, pi. 55, figs E, F. 'Indes Orientales' [? Ambon] (type-material 

lost). 
Papilio isse Cramer; Stoll, 1781 : 95, pi. 339, figs C, D. 'Amboina'. 
Papilio Danai candidi sp., Zschach, 1788 : 88, no. 31*. 'Exoticus'. 
Papilio bicolor Gmelin, 1790 : 2261, no. 880. LECTOTYPE <$, 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy by Kirby, 1869 : 357. 
Papilio bicolor; O'Reilly, 1813 : 74, no. 31. 'India'. 
Pieris isse (Cramer) ; Kirby, 1869 : 357. 

Delias isse (Cramer); Kirby, 1871 : 476; Kirby, 1880 : 321; Talbot, 1929 : 88. 
Delias isse isse (Cramer); Talbot, 1937 : 473; D'Abrera, 1971 : 148. 

Papilio bicolor Gmelin. A single male specimen in Dublin bears the labels 
/31/Isse, Cram. (31) (Bicolor, Gmel, type specimen)/, the latter in Kirby's hand- 
writing on blue paper; this specimen is hereby designated lectotype of Papilio 
bicolor Gmelin, and has been labelled accordingly. It lacks the antennae and 
abdomen, but is otherwise in fair condition. The lectotype fits the description 
given by Talbot (1937 : 473) for Delias isse isse (Cramer), known from Ambon, 
Ceram, Gisser and Banda. Both isse and bicolor were most probably obtained 
from Ambon. 

Papilio isse Cramer. A male of this species in the BMNH, ex Felder collection, 
bears a 'Cramer label' (see p. 56) for isse. This specimen does not, however, 
correspond as closely to the original Cramer figure as two other ex Felder specimens 
which bear no other data. The BMNH does not appear to have the types of Papilio 



32 R. I. VANE-WRIGHT 

crocale or P. celmus, described on the same page as isse, all from specimens loaned 
to Cramer by B. Vriends of Haarlem (Cramer, 1775 : 87). It would appear possible 
that the 'Cramer label' was attached to a fresh specimen obtained at a later date. 
Stoll (1781) described the female of isse from one or more specimens collected on 
Ambon, in the collection of Baron Rengers. The BMNH collections include several 
female specimens of isse, ex Felder, which bear no further data. It is probable 
that some of these, and the males noted above, originated from the Rengers collec- 
tion, or came from the same source. 



Delias ceneus ceneus (Linnaeus) [Pieridae] 

Papilio ceneus Linnaeus, 1758 : 487. LECTOTYPE 9- ' m Indiis' [Ambon] (MLU), here 

designated. Correct type-locality : Corbet, 1949 : 195. 
[Papilio hyparete Linnaeus ; Clerck, 1 764 : pi. 38, two figures at lower margin. Misidentification.T 
Papilio hyparete Linnaeus, varietas an Femina?; Linnaeus, 1764 : 247. 
Papilio caeneus Linnaeus; Linnaeus, 1767 : 766. 
[Papilio hyparete Linnaeus; Cramer, [1779] : 30, pi. 210, figs A, B. 'Amboina'. Misidenti- 

fication.] 
[Papilio hyparete Linnaeus; Stoll, [1781] 195-6, pi. 339, figs E, F. 'Amboina'. Misidenti- 

cation.] 
Papilio Danai candidi sp., Zschach, 1788 : 88, no. 32.* 'Exoticus'. 
Papilio discors Gmelin, 1790 : 2261, no. 881. LECTOTYPE $, 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy by Kirby, 1871 : 476. 
[Papilio antonoe Cramer; Herbst, 1792 : 126, pi. 100, figs 1, 2, 3, 4. Misidentification.] 
Papilio plexaris Donovan, 1805 : pi. 18, two figures. 'Botany Bay' [? Ambon] (type-material 

lost). Synonymy from Talbot, 1937 : 544. 
Papilio discors; O'Reilly, 1813 : 74, no. 32. 'India'. 
Pieris philyra Godart, 1819 : 150. LECTOTYPE 9> [Ambon] (MLU), here designated. 

Synonymy from Talbot, 1937 : 544- 
Cathaemia anthyparete Hiibner, [1819] : 92. Ambon (type-material lost). Synonymy 

from Talbot, 1937 : 544. 
Pieris discors (Gmelin); Kirby, 1869 : 357. 

Delias caeneus (Linnaeus); Kirby, 1871 : 476; Kirby, 1880 : 321; Talbot, 1929 : 88. 
Delias caeneus caeneus (Linnaeus); Talbot, 1937 : 543- 
Delias ceneus ceneus (Linnaeus); Corbet, 1949 : 195. 
Delias caeneus caeneus (Linnaeus); D'Abrera, 1971 : 150. 

Papilio discors Gmelin. Two specimens, a pair, exist in the NMI collection; 
they appear above the drawer label /Types of Discors, Gmel./. One specimen has 
no further data, while the other simply bears the addition j^2j. The latter specimen, 
the male, is hereby designated lectotype of Papilio discors Gmelin, and has been 
labelled accordingly. The female (the 'altero sexu' of the original description) 
is similarly designated paralectotype; it lacks the abdomen. The lectotype is in 
fair condition, but as with most of the Leske material, is now very faded. Probably 
from Ambon, discors fits the description given by Talbot (1937 : 544), and is thus 
best treated as strictly synonymous with ceneus ceneus (Linnaeus) . It is interesting 
to note that Zschach recognized the sexual dimorphism of this species. 

Papilio ceneus Linnaeus. Corbet (1949 : 195) summarizes his work on the 
type-material of this species. I hereby designate the specimen in MLU as dealt 



BUTTERFLIES NAMED BY GMELIN 33 

with by Corbet (loc. cit.) and illustrated by Clerck (1764 : pi. 38, two lowermost 
figures, as Papilio hyparete, in part), lectotype of Papilio cenens Linnaeus. 

Papilio plexaris Donovan was described from at least two male specimens from 
'Botany Bay'. Donovan's natural history collection was auctioned by Stevens 
in 1818 (Horn & Kahle, 1935 : 60); the types are presumed to be lost. 

Pieris philyra Godart was described by reference to Cramer's (1779), Stoll's 
(1781) and Herbst's (1792) figures of 'hyparete', and also, apparently, from material 
actually studied by Godart. In addition, Godart gives a first reference to 'Papilio 
Hyparete. Variet. anfem.P Linn. Mus. Lud. Ulr. p. 247.'. As summarised by Corbet 
(1949), this last reference can be traced to the lectotype of Papilio ceneus Linnaeus, 
as designated above. To confirm existing synonymy, I hereby designate the 
lectotype specimen of Papilio ceneus Linnaeus as, additionally, the lectotype of 
Pieris philyra Godart, of which the latter thereby becomes an objective synonym. 
The specimens represented by the illustrations of Cramer (1779 : pi. 210, figs A, 
B), Stoll (1781 : pi. 339, figs E, F) and Herbst (1792 : pi. 101, figs 3, 4, 5) are desig- 
nated paralectotypes. Of these it is interesting to note that while the figures of 
Cramer and Stoll, under the misidentified name hyparete, do actually refer to ceneus 
( = philyra), the figures of Herbst actually refer to true hyparete Linnaeus. Herbst 
did, however, illustrate ceneus on the previous plate (pi. 100, figs 1, 2, 3, 4), this 
time under the misidentified name antonoe Cramer. A pair of specimens in the 
MNHN have been recognized as paralectotypes of Pieris philyra, and labelled 
accordingly. A male ceneus in the Rothschild collection, BMNH, bearing the 
'Cramer label' /No. 32. hyparete. Cr. Ill 210, A.B. & IV 339, CD./, is probably 
a van Lennep specimen as referred to by Cramer (1779 : 30), but is evidently not 
the one figured by Cramer on plate 210, and has not therefore been designated as 
a paralectotype. 

Cathaemia anthyparete Hubner was named by reference to 'Hyparete Cram. 210 
A.B., & 339 E.F.'. The figures on plate 339 (Stoll, 1781) are from one or more 
female specimens received from Ambon by Baron Rengers (Stoll, op. cit.: 96); 
the material is apparently lost or unrecognizable. 

The name Papilio ceneus Linnaeus might be regarded as a senior primary homonym 
of the name Papilio cenea Stoll, as published in 1790 in the Supplement to 'Cramer'. 
The name cenea Stoll is in current usage for the southern African subspecies of 
Papilio dardanus Brown. A large volume of very important genetical and other 
biological work has been done on the races of this polymorphic mimic species, 
and a strong case could be made out for conserving the junior homonym, Papilio 
dardanus cenea Stoll. Fortunately, as Charles Cowan has pointed out to me, 
Ceneus is a girl's name, while 'cenea' is not a Latin word but an arbitrary combina- 
tion of letters. Therefore no question of homonymy arises, and both names can 
stand. 

Delias dorimene (Stoll) [Pieridae] 

Papilio dorimene Stoll, [1782] : 201, pi. 387, figs C, D. Holotype g, Ambon (type-material 

lost). Correct author and date: Hemming, 1958 : 43. 
Papilio Danai candidi sp., Zschach, 1788 : 88, no. 34*. 'Exoticus'. 



34 R- I. VANE-WRIGHT 

Papilio fuliginosus Gmelin, 1790 : 2261, no. 882. LECTOTYPE^, 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy by Kirby, 1869 : 357. 
Papilio dorimene Cramer; Herbst, 1792 : 132, pi. 102, figs 6, 7. 
Papilio fuliginosus; O'Reilly, 1813 : 74, no. 34. 'India'. 
Pieris ageleis Godart, 1819 : 113, 147. Ambon (type-material lost). Synonymy from Talbot, 

1937 = 447- 
Pieris dorimene (Cramer); Kirby, 1869 : 357. 
Delias dorimene (Cramer); Kirby, 1871 : 476. 
Delias dorimene avenda Fruhstorfer, 1912& : 5. LECTOTYPE $, Ceram (BMNH), here 

designated [examined]. Synonymy from Talbot, 1937 : 447. 
Delias dorimene (Cramer); Talbot, 1929 : 88; Talbot, 1937 : 447, 587; D'Abrera, 1971 : 146. 

Papilio fuliginosus Gmelin. A single male specimen in Dublin bears the labels 
/34/ Type of Fuliginosus Gmel./; this specimen is hereby designated lectotype of 
Papilio fuliginosus Gmelin, and has been labelled accordingly. The specimen 
lacks the abdomen, but is otherwise in fair condition. The lectotype probably 
came from Ambon; it fits the description given by Talbot (1937), who did not 
recognize any subspecies of dorimene, which is found on the S. Moluccan islands 
of Ceram, Gisser, Saparua and Ambon. As also noted by Talbot (op. cit.: 587), 
fuliginosus (Gmelin) preoccupies Delias fuliginosus Kenrick, 1909. 

Papilio dorimene Stoll was apparently described from a single male specimen, 
received from Ambon by Raye de Breukelerwaert. This specimen, the holotype, 
illustrated in Stoll's original figures, appears to be lost; no certain Stoll or Cramer 
specimens of this taxon exist in the Rothschild collection (BMNH), or in Leiden 
(RNH), where such might have been expected. 

Pieris ageleis Godart was described by reference to the original description of 
'Papillon dorimene Cramer', figures in Herbst (1792), and specimens examined by 
Godart himself. None of Godart's specimens can be traced in Paris or Edinburgh 
and, as noted above, the specimen illustrated by Stoll (1782 : pi. 201, figs C, D) 
is also apparently lost or destroyed. 

Delias dorimene avenda Fruhstorfer was described from a pair of specimens 
in the Fruhstorfer collection from 'Ceram', the male collected by Ribbe, the female 
by Kuhn. A single male now in the BMNH bears the labels /Type/Ceram Jllo 
C. Ribbe 1884/dorimene avenda Frhst [in Fruhstorfer's handwriting] /Fruhstorfer 
Coll BM 1937-285/; this specimen is hereby designated lectotype of Delias dorimene 
avenda Fruhstorfer, and has been labelled accordingly (type no. Rh 17308). A 
single female bearing the labels /Type/O. Ceram Fruhstorfer/avenda Fruhst. $ 
[in Talbot's handwriting] /Frushtorfer Coll BM 1937-285/, is similarly designated 
paralectotype. 



Eurema hecabe hecabe (Linnaeus) [Pieridae] 

Papilio hecabe Linnaeus, 1758 :47c. LECTOTYPE $, 'in Asia' [China: Canton] (LC), here 
designated [examined]. Correct type-locality: Corbet, 1941&; 1942; 1949. 

Papilio Danai candidi sp., Zschach, 1788 : 89, no. 35*. 'Exoticus'. 

Papilio chrysopterus Gmelin, 1790 : 2261, no. 883. LECTOTYPE £, 'extra Europam' 
[? S. E. China] (NMI), here designated [examined]. Syn. n. 

Papilio chrysopterus; O'Reilly, 1813 : 74, no. 35. 'India'. 



BUTTERFLIES NAMED BY GMELIN 35 

Papilio chrysopterus Gmelin; Kirby, 1869 : 357. 
Eurema chrysopterus (Gmelin); Kirby, 1871 : 450. 
Terias blanda acandra Fruhstorfer, 1910 : 169. Holotype $, Hong Kong (BMNH), [examined]. 

Joicey & Talbot, 1924 : 533; synonymy from Talbot, 1935 : 553- 
Papilio chrysopterus Gmelin; Talbot, 1935 : 622. 
Papilio hecabe Linnaeus; Corbet, 19416 : 27. 
Eurema hecabe hecabe (Linnaeus) ; Corbet, 1949 : 494. 

Papilio chrysopterus Gmelin. A single male specimen in Dublin bears the labels 
/Type of Chrysopterus, Gmel. [in Kirby's handwriting, on blue paper] j$2 : 1925/ 
Examined by H. T. G. Watkins in 1925 and pronounced to be synonymous with 
Terias hecabe hecabe (Linne) from S. China. $j. This specimen is hereby designated 
lectotype of Papilio chrysopterus Gmelin, and has been labelled accordingly; al- 
though lacking both antennae, it is otherwise in fair condition. 

H. T. G. Watkins, in collaboration with N. D. Riley, evidently closely compared 
the lectotype of chrysopterus with material of hecabe in the BMNH; a male hecabe 
from Hainan is labelled j chrysopterus, Gmel. compared with type 12. v. 1925. N.D.R., 
H.W./i and a specimen from Macao, S. China, is similarly labelled /probably nearest 
to hecabe type/. All three specimens are virtually identical. However, the Hainan 
population is currently dealt with as a separate subspecies, E. hecabe subdecorata 
(Moore) (Corbet & Pendlebury, 1932 : 160). It would seem much more likely 
that Leske's material would have come from one of the trading ports on the China 
coast, than from the little known island of Hainan. The type-locality of hecabe 
has also been the subject of doubt. Corbet (1941b; 1942; 1945; 1949) has shown 
it to be Canton; I therefore propose to regard Canton as the type-locality of chrysop- 
terus Gmelin, which thereby becomes a strict synonym of Eurema hecabe hecabe 
(Linnaeus). 

Despite the fact than many of the Leske type-specimens evidently came from 
Ambon, in this case it is clear, from examination of specimens, that chrysopterus 
does not represent the race of hecabe from that island (E. h. diver sa (Wallace)). 

Papilio hecabe Linnaeus. Corbet has studied the type-material of this species 
in considerable detail. It remains only to provide a formal lectotype designation. 
I hereby designate the female specimen in the Linnaean Collection, London, which 
bears the labels /Hecabe 763/74. Hecabe/, as lectotype of Papilio hecabe Linnaeus. 
The lectotype is illustrated by Corbet & Pendlebury (1956 : pi. 29, fig. 6). 



Euploea phaenareta phaenareta (Schaller) [Danaidae] 

Papilio sp., Seba, 1765 : 25, pi. 19, figs 13, 14. 

[Papilio midamus Linnaeus; Cramer, [1780] : 131-2, pi. 266, figs A, B. 'Amboina'. Misidenti- 

fication.] 
Papilio phaenareta Schaller, 1785 : 177, pi. 5, figs 1, 2. [? Ambon] (type-material destroyed). 
Papilio Danai festivi sp., Zschach, 1788 : 89, no. 46*. 'Exoticus'. 
Papilio affinis Gmelin, 1790 : 2289, no. 885. LECTOTYPE 9. 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy by Kirby, 1869 : 358. [Junior primary 

homonym of Papilio affinis Fabricius, 1775.] 
[Papilio midamus Linnaeus; Herbst, 1793 : 12, pi. 119, figs 1, 2. Misidentification.] 
Papilio affinis] O'Reilly, 1813 : 74, no. 46. 'India'. 



36 R. I. VANE-WRIGHT 

Trepsichrois alea Hiibner, 1816 : 16. Ambon (type-material lost). Synonymy from 

Bryk, 1937 : 370. 
Danais prothoe Godart, 1819 : 177. LECTOTYPE g, Ambon (RSM), here designated 

[examined]. Synonymy from Grimshaw, 1898 : 2. 
Euploea phaenareta (Schaller) ; Kirby, 1869 : 358. 
^Euploea affinis (Gmelin) ; Kirby, 1871 : 17. 
Euploea phaenareta (Schaller); Kirby, 1871 : 639. 
Euploea phaenareta phaenareta (Schaller); Bryk, 1937 : 369; D'Abrera, 1971 : 190. 

Papilio affinis Gmelin. A single female specimen in Dublin bears the label 46 
Phaenareta, Schall. type of Affinis, Gmel./, in Kirby's handwriting. This specimen 
is hereby designated lectotype of Papilio affinis Gmelin, and has been labelled 
accordingly. This specimen which, apart from lacking the abdomen, is in fair 
condition, corresponds closely with material of the nominate subspecies of Euploea 
phaenareta from Ceram and Ambon (D'Abrera, 1971 : 190), probably coming, in 
fact, from the latter island. The 'sexus alter' of the original description appears 
to have been destroyed; no other specimens of phaenareta are now present in the 
NMI. The specific name affinis Gmelin is in any case invalid, being a junior primary 
homonym of Papilio affinis Fabricius. 

Papilio phaenareta Schaller was described from at least one male specimen studied 
by Schaller, and by reference to female specimens illustrated by Seba (1765) and 
Cramer (1780), the last with some reservation. Schaller's material most probably 
came from Ambon, as with most other contemporary material from the Moluccas. 
The Schaller collection has been destroyed (Horn & Kahle, 1936 : 241). 

Trepsichrois alea Hiibner was described by reference to 'rnidamus Cram. 266. 
A.B.'. Cramer's material (1780 : 132, pi. 266, figs A, B) came from Ambon. An 
ex Felder specimen of phaenareta in the Rothschild collection (BMNH), which 
bears the 'Cramer label' /No. 62. Midamus. Cr.III.266.A.B./ and a Felder /Coll. 
Lenep/ label, might be the specimen originally illustrated. However, I have com- 
pared this specimen with Cramer's original pattern plate, but find the finer details, 
especially on the underside, just too different to be confident of such an identifica- 
tion. 

Danais prothoe Godart was described by reference to the misidentified illustrations 
of 'midamus'' given by Cramer (1780) and Herbst (1793), in addition to one or more 
specimens studied by Godart himself. A single male specimen now in the RSM, 
ex Dufresne collection, bears the label /Amboine/ (Grimshaw, 1898 : 2). This 
specimen, which I have examined in Edinburgh, is hereby designated lectotype of 
Danais prothoe Godart. 



Danaus limniace exoticus (Gmelin) stat. n., nom. rev. [Danaidae] 

(PL 6, figs 21, 22) 

Papilio Danai festivi sp., Zschach, 1788 : 89, no. 46b*. 'Exoticus'. 

Papilio exoticus Gmelin, 1790 : 2289, no. 886. LECTOTYPE 3\ 'extra Europam' [? Ceylon 7 

(NMI), here designated [examined]. 
Papilio exoticus; O'Reilly, 1813 : 74, no. 46b. 'India'. 



BUTTERFLIES NAMED BY GMELIN 37 

Danais leopardus Butler, 1866 : 52. LECTOTYPE <J, India (BMNH), here designated 

[examined]. Syn. n. 
Danaus limniace (Cramer); Kirby, 1869 : 358. 
?Danus exoticul [sic] (Gmelin); Kirby, 1871 : 8. 
Danaus limniace (Cramer) ; Kirby, 1871 : 639. 
Danaida limniace mutina Fruhstorfer, 1910 : 204, pi. 78, row a. LECTOTYPE $, Ceylon 

(BMNH), here designated [examined]. Syn. n. 
Danaus limniace leopardus (Butler); Talbot, 1943 : 134; Talbot, 1949 : 31. 
Danaus limniace mutina (Fruhstorfer); Woodhouse, 1950 : 29. 
Danais limniace (Cramer); Wynter-Blyth, 195 1 : 67. 
Danaus limniace leopardus (Butler); Bailey, 1951 : 74; Corbet & Pendlebury, 1956 : 440. 

Papilio exoticus Gmelin. A single male specimen in Dublin bears the label 
/46b Zschach. Probably Leskean type of P.exoticus Gmel./, in Kirby's handwriting, 
on blue paper. Despite Kirby's doubt, the original description by Zschach of the 
male scent organ ('squama subtus') of this species, makes it fairly certain that this 
specimen was part of the type-material of exoticus. This specimen is, therefore, 
hereby designated lectotype of Papilio exoticus Gmelin, and has been labelled 
accordingly. It is generally in fair condition, but lacks the head. 

P. exoticus fits the diagnosis given by Talbot (1943 : 134) for Danaus limniace 
leopardus (Butler), and is very similar to many specimens in the BMNH of this 
subspecies, from Ceylon and India. It clearly does not belong to any of the other 
limniace subspecies enumerated by Talbot, both from examination of the BMNH 
collection, and study of Talbot's Revisional Notes. At the subspecies level this 
then places leopardus Butler and mutina Fruhstorfer in the synonymy of exoticus; 
lectotype designations for these taxa are appended below. It should be noted 
that the lectotypes of exoticus and mutina (PI. 6, fig. 21) agree well with each other, 
and with the great majority of specimens from the Indian region dealt with by 
Talbot as leopardus. All three type-specimens of leopardus (PI. 6, fig. 22), however, 
are atypical of the Indian subspecies, not showing any fusion of the fore wing 
posterior stripes or distal spot (Talbot, 1943), and thus not actually fitting Talbot's 
diagnosis for subspecies leopardus. However, the types appear to fall within the 
range of variation of the Indian subspecies, and so I am placing all three names, 
exoticus, leopardus and mutina, in synonymy. If further subdivision ever proves 
necessary, it would appear best to consider the lectotype of exoticus as being from 
Ceylon. 

The name limniace leopardus (Butler) might have qualified for conservation 
under the reworded Article 23b of the International Code as advocated by Mayr 
et al. (1971), but this was rejected by the Zoological Congress held at Monaco. 
Further, as the type-specimens of leopardus are atypical of the Indian region sub- 
species, and the three most recent major books on Indian region butterflies use 
different names (Talbot, 1949; Woodhouse, 1950; Wynter-Blyth, 1951), it seems 
most straightforward to adhere to priority. 

Danais leopardus Butler was described from three syntypic specimens, from 
'India'. All three are extant in the BMNH, comprising one male and two females. 
The male bears the following labels: /Type [red] /N.India 43.10/Tirumala leopardus 
Butler Type/B.M.Type no Rh.6417 Danais leopardus $ Butl./. This specimen 



38 R. I. VANE-WRIGHT 

is hereby designated lectotype of Danais leopardus Butler (PI. 6, fig. 22), and has 
been labelled accordingly. The females, /Punjab 54.74/Type no.6418/ and /Moul- 
mein 43.43/Type no. 6419/ respectively, are similarly designated paralectotypes. 

Danaida limniace mutina Fruhstorfer was described from an unstated number 
of specimens, from 'Ceylon'. The BMNH possesses five males and four females 
from Ceylon which were received in the Fruhstorfer Danaid collection. One male 
bears the following labels: /Ceylon 1889 H.Fruhstorfer/Type [red; Fruhstorfer 
pattern] /limniace mutina Fruhst. [in Fruhstorfer 's handwriting] /Fruhstorfer Coll. 
BM 1937-285/. This specimen is hereby designated lectotype of Danaida limniace 
mutina Fruhstorfer (PI. 6, fig. 21), and has been labelled accordingly (type no. 
Rh 17296). The remaining four males and four females are similarly designated 
paralectotypes (3^, 3 $, /Ceylon 1889 H. Fruhstorfer'; 1 ^, /Slid Ceylon Mai 1889 
H. Fruhstorfer/; 1 $, Ceylon Schilling ex Coll. H. Fruhstorfer/). 



Danaus juventa claviger (Gmelin) stat. n., nom. rev. [Danaidae] 

(PI. 6, fig. 20) 

Papilio Danai festivi sp., Zschach, 1788 : 89, no. 46c*. 'Exoticus'. 

Papilio claviger Gmelin, 1790 : 2289, no. 887. LECTOTYPE 3\ 'extra Europam' [? Ambon] 
(NMI), here designated [examined]. 

Papilio claviger; O'Reilly, 1813 : 75, no. 46c. 'India'. 

Danais meganire Godart, 1819 : 192. LECTOTYPE <$, 'Java' [? Ambon] (MNHN), here 
designated [examined]. Synonymy originally given by Kirby, 1871 : 639. 

Papilio claviger Gmelin; Kirby, 1869 : 358. 

IDanaus claviger (Gmelin); Kirby, 1871 : 8. 

Danaus claviger (Gmelin); Kirby, 1871 : 639; Kirby, 1877 : 691; Ribbe, 1889 : 218, 'Amboina 
und Ceram'; Rober, 1891 : 289, 'Ceram'. 

Radena buruensis Holland, 1900 : 56. LECTOTYPE g, Buru (CM), here designated [photo- 
graph examined]. Synonymy with meganire: Talbot, 1943 : 144. Syn. n. 

Wanaida juventa ogylla Fruhstorfer, 1910 : 214. LECTOTYPE £, Aru (BMNH), here 
designated [examined], Synonymized with meganire by D'Abrera, 1971 : 172. 

Danaus juventa meganire (Godart); Bryk, 1937 : 167; Talbot, 1943 : 144; D'Abrera, 1971 : i; 2 - 

Papilio claviger Gmelin. A single male specimen in Dublin bears the labels 
/46c/Claviger, Gmel. Leskean type 11.460?/, in Kirby 's handwriting. This specimen 
is hereby designated lectotype of Papilio claviger Gmelin, and has been labelled 
accordingly; it is in fair condition, but lacks the abdomen (a microscope slide of 
one foreleg has been prepared to check the sex.). 

P. claviger fits very well with the material dealt with by Talbot (1943 : 144) 
as Danaus juventa meganire (Godart), from Buru, Ceram, Gisser, Nausa Laut 
and Ambon, probably coming from the last named island. There seems no doubt that 
claviger is the senior name for the race of juventa on this group of islands, as accepted 
by Ribbe (1889 : 218) and Rober (1891 : 289). 

The original type-locality of meganire was given by Godart (1819 : 192) as Java: 
all subsequent treatments seem to follow Boisduval (1832 : 105) and Blanchard 
( I 853 : 387-388) in attributing Godart's species to the S. Moluccas group of islands. 



BUTTERFLIES NAMED BY GMELIN 39 

I have been able to confirm this by examination of the lectotype (PI. 6, fig. 20), 
designated below, which probably came from Ambon. 

D'Abrera (1971 : 172) has relegated the name ogylla Fruhstorfer, as dealt with 
by Talbot (1943 : 144), to the formal synonymy of meganire (— claviger). Material 
formerly in the Rothschild collection, evidently not examined either by Talbot 
or D'Abrera, does not help to make an easy decision on this; it does appear that 
juventa from the small islands between Ceram and Kai have a somewhat different 
appearance, as discussed by Talbot under ogylla, but probably the effect is a clinal one. 

Danais meganire Godart was described from an unstated number of specimens, 
from 'Java'. A single male specimen in Paris bears the labels /Type/Meganira 
God. Indes orient, [yellow paper] /Museum de Paris/. This specimen is hereby 
designated lectotype of Danais meganire Godart (PL 6, fig. 20), and has been labelled 
accordingly. The lectotype lacks antennae and abdomen; in addition the fore 
legs are embedded in glue. The sex has been determined from the colour and 
scaling of the hind wing anal cells. A second male specimen in Paris which has 
the labels /Type/Indes orientales. Museum de Paris/, and the two specimens in 
the RSM, noted by Grimshaw (1898 : 2), are similarly designated paralectotypes. 

Radena buruensis Holland was described from 15 examples collected on Buru 
by William Doherty. These specimens (six males, nine females) are all extant 
in the Carnegie Museum, Pittsburgh. One male, which bears the labels/Radena 
Bouruensis Holland n.sp./[in Holland's handwriting] /Bourou. Coll. Doherty/Holland 
Collection/, is hereby designated lectotype of Radena buruensis Holland, and has 
been labelled accordingly (by Dr Clench, on my behalf). I have examined a photo- 
graph of this specimen, kindly taken by Dr F. Martin Brown. The remaining 
14 specimens are similarly designated paralectotypes. 

Danaida juventa ogylla Fruhstorfer was described from an unstated number of 
specimens from Aru. A single male in the BMNH bears the labels /Aru-Jnsel/ 
Type/ogylla Fruhst. [in Fruhstorfer's handwriting]/ Fruhstrofer Coll. BM 1937- 
285/. This specimen is hereby designated lectotype of Danaida juventa ogylla 
Fruhstorfer and has been labelled accordingly (type no. Rh 17304). 



Taenaris urania urania (Linnaeus) [Amathusiidae] 

Papilio urania Linnaeus, 1758 : 466. LECTOTYPE $, 'in Indiis' [Ambon] (MLU), here 

designated. Correct type-locality: Corbet, 1949 : 198. 
[Papilio cassiae Linnaeus; Clerck, 1764 : pi. 29, fig. 3. Misidentification; identity with urania 

established by Corbet, 1941a : 15; Corbet, 1949 : 198.] 
Papilio jairus Cramer, 1775 : 9, pi. 6, figs A, B. 'East Indies' [? Ambon] (specimen lost?). 

Syn. n. 
Papilio jairus Cramer; Cramer, [1777] : 134-135, pi. 185, figs A, B, C. 'Amboina'. 
Papilio Danai festivi sp., Zschach, 1788 : 89, no. 47*. 'Exoticus'. 
Papilio mavinus Gmelin, 1790 : 2289, no. 888. LECTOTYPE 3\ 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy by Kirby, 1869 : 358. 
Oreas dubia Jaira Hiibner, [1808] : pi. [84], figs 1,2. [? Ambon] (specimen lost?). Correct 

date: Hemming, 1937a : 336, 403. [Junior secondary homonym of jairus (Cramer), 1775.] 

Syn. n. 
Papilio murinus [sic]; O'Reilly, 1813 : 75, no. 47. 'India'. 



4 o R. I. VANE-WRIGHT 

Tenaris nysa Hubner, [1819] : 53. Lectotype <$, Ambon (BMNH), designated by Hemming. 

1964 : 94 [examined: see below; see also Hemming, 1967 : 426]. Synonymy from Kirby, 

1871 : 117. 
Drusilla urania (Linnaeus); Kirby, 1869 : 358. 
Tenaris urania (Linnaeus); Kirby, 1871 : 117; Kirby, 1880 : 300. 
Tenaris nox Kirby, [1908] : 47 [new name for Papilio jaira Hubner ]. Syn. n. 
Taenaris urania urania (Linnaeus) ; Stichel, 1933 : 88. 
Tenaris urania urania (Linnaeus); Brooks, 1950 : 232. 
Taenaris urania urania (Linnaeus); D'Abrera, 1971 : 292. 'Ambon, Saparua'. 

Papilio marinus Gmelin. A single male in Dublin bears the labels /47?/47 
Urania ? type of Marinus, Gmel./, in Kirby's handwriting, on blue paper. This 
specimen is hereby designated lectotype of Papilio marinus Gmelin, and has been 
labelled accordingly. It is in fair condition. 

Brooks (1950), in his revision of Taenaris, does not mention marinus, but in 
his arrangement of the BMNH Amathusiid collection he included it as a strict 
synonym of urania, the type-locality of which is now regarded as Ambon (Corbet, 
1949). The three subspecies of urania currently recognized (Brooks, op. cit.; 
D'Abrera, 1971) are not, however, readily separable on the basis of single specimens, 
but it seems likely that all the old types, including that of marinus, came from 
Ambon. As noted by Kirby (1869 : 358) and also evidently by O'Reilly (1813 : 75), 
the name marinus is really a lapsus by Gmelin for murinus, which word is used 
in the original description. However, since the taxon in question is almost certainly 
relegated to permanent synonymy, the original spelling of Gmelin is best retained. 

The names jairus Cramer, nox Kirby ( = jaira Hubner) and nysa Hubner, listed 
in the synonymy above, have been dealt with for a long period in the literature 
as forms of Taenaris urania urania (Brooks, 1950; Hemming, 1964 : 123). However, 
all these names are available, having been proposed as full species, and as such 
should be listed in the synonymy of urania (L.). The present International Code 
does not apply to infrasubspecific terminology, but these names may still be applied 
to the dubious forms of urania (not, in my opinion, a truly polymorphic species) 
by those who may wish to do so. It should be noted that the name duplex Stichel, 
referred to by Brooks (1950 : 232) as a form of urania urania, correctly applies 
to Taenaris urania pandemos Fruhstorfer. 

During this work I have recognized the lectotype-specimen of Tenaris nysa 
Hubner in the BMNH collections. Hemming (1964 : 94) made the original designa- 
tion on the basis of a figure in Cramer (1777), without realising that some of the 
Cramer material was still in existence (as discussed elsewhere in this paper). The 
discovery of this, lectotype-specimen is of special significance, nysa beirg the type- 
species of the genus Taenaris Hubner (Hemming, 1967 : 426). 

Tenaris nysa Hubner. Lectotype designated by Hemming (1964 : 94) as the 
specimen represented by figure A on plate 185, in Cramer (1777). This specimen, 
a male, is now believed to be in the BMNH, and bears the labels /No. 94. Jairus. 
Cr. I. 6. AB. & II. 185. ABC/Felder Colin/Rothschild Bequest BM 1 939-1 /Tenaris 
nysa Hubner, Lectotype $, det. R.I.V.-W., 1972/BM Type No. Rh 17305/. The 
first label is a typical 'Cramer label'; Cramer (1777 : 135) notes the specimens 
illustrated as being from Ambon, in the collection of C. van Lennep. 



BUTTERFLIES NAMED BY GMELIN 41 

Papilio urania Linnaeus. Corbet (1949 : 198) summarizes his work on the 
type-material of this species. I hereby designate the specimen in MLU, as dealt 
with by Corbet (loc. cit.), and illustrated by Clerck (1764 : pi. 29, fig. 3) under 
the name Papilio cassiae, lectotype of Papilio urania Linnaeus. 

Papilio jairus Cramer was originally described from an unstated number of 
females, from the 'East Indies'. Those parts of the Cramer material which have 
not survived in the Felder collection or at Leiden are presumed to be lost. I have 
been unable to find any urania specimens of the distinctive nature of the specimen 
described as jairus by Cramer in the Felder material in the BMNH, and likewise 
no specimens of this form appear to survive in Leiden. 

Oreas dubia jaira Hiibner was based on one or more male specimens, as originally 
illustrated by Hiibner. The locality was not indicated, and no published or manu- 
script text exists for the particular plate concerned (Hemming, 1937a; 19376)- 
The name jaira Hiibner is invalid, being a secondary homonym of jairus Cramer; 
Kirby proposed the name Tenaris nox to replace it. 



Euploea leucostictos leucostictos (Gmelin) [Danaidae] 

Papilio Danai festivi sp., Zschach, 1788 : 90, no. 48*. 'Exoticus'. 

Papilio leucostictos Gmelin, 1790 : 2289, no. 889. LECTOTYPE [$], 'extra Europam' [Ambon] 
(NMI), here designated [examined]. Corbet, 1947 : pi. 4, fig. 1. 

Lemnas mutabilis Nemertes Hiibner, [1807] : pi. [26], figs 3, 4; manuscript text, Hemming, 
19376 : 102-104. [Ambon] (specimen lost). Synonymy by Kirby, 1869 : 359; Corbet, 
1947 : 229. 

Papilio leucostictos; O'Reilly, 1813 : 75, no. 48. 'India'. 

Euplaea aglidice Boisduval, 1832 : 96. LECTOTYPE $, 'Rawack' [Ambon] (BMNH), here 
designated [examined]. Syn. n. 

Euploea pasithea Felder & Felder, 1865 : 318. LECTOTYPE <J, Ambon (BMNH), here desig- 
nated [examined]. Synonymy by Kirby, 1871 : 13. 

Euploea leucostictos (Gmelin); Kirby, 1869 : 358; Kirby, 1871 : 13; Kirby, 1880 : 294. 

Salpinx oculata Moore, 1883 : 302. LECTOTYPE <^, 'Mindanao' [Ambon] (BMNH), here 
designated [examined]. Syn. n. 

Euploea leucostictos leucostictos (Gmelin); Corbet, 1947 : 228, pi. 4, fig. 1. (Correct type- 
locality). 

Euploea nemertes nemertes (Hiibner); D'Abrera, 1971 : 192. 

Papilio leucostictos Gmelin. A single, composite, specimen in Dublin bears the 
labels /48/48 Leucostictos, Gmel. Type from Mus.Lesk./, the latter being in Kirby's 
handwriting on blue paper. This specimen (figured by Corbet, 1947 : pi- 4, fig- 1) 
is hereby designated lectotype of Papilio leucostictos Gmelin, and has been labelled 
accordingly. The wings are those of a female insect, but the abdomen is male; 
the head is glued on upside down with two bristles affixed as antennae. The latter 
deception was noted by Corbet and clearly shows in his photograph. Corbet (op. 
cit.) also established the true type-locality of leucostictos, thereby correcting the 
erroneous locality of Java which had been attributed to this name for many years 
(e.g. Bryk, 1937 : 392). As a result, the names Lemnas nemertes Hiibner, Euplaea 
aglidice Boisduval, Euploea pasithea Felder & Felder and Salpinx oculata Moore 



42 R. I. VANE-WRIGHT 

fall into the strict synonymy of leucostictos Gmelin. Lectotype designations for 
three of the synonymous species are appended. 

Lemnas nemertes Hiibner was described from an unstated number of male speci- 
mens, now lost. Hemming (1937a : 419, 429; 1937& : 102-104) deals with the 
previously unpublished manuscript text; the original material evidently came 
from Ambon. 

Euplaea aglidice Boisduval was described from an unstated number of female 
specimens, from 'Rawack (terre de Papous)'. A single female in the BMNH bears 
the following labels: /Type/Typicum Specimen/Aglidice Boisduval Astrolabe - type/ 
Ex Musaeo Dr Boisduval/vu par Moore en 1 881 /Euplaea aglidice Bdv (astrolabe, 
page 96) $ Rawack (terre de Papous) /Euploea nemertes. $. Hubn./Ex Oberthiir 
Coll BM 1927-3/. This specimen is hereby designated lectotype of Euplaea aglidice 
Boisduval, and has been labelled accordingly (type no. Rh 17297). 

Euploea pasithea Felder & Felder was described from an unstated number of 
male and female specimens, from 'Amboina', collected by Doleschall and Lorquin. 
The BMNH possesses a single pair, the male of which bears the following labels: 
/Amboina Doleschall/Felder Colin/Type HT/pasithea/Rothschild Bequest BM 
1939-1/. This specimen is hereby designated lectotype of Euploea pasithea Felder 
& Felder, and has been labelled accordingly (type no. Rh 17298). The female 
specimen bears the labels /Amboina Lorquin/Pasithea de Haan/Felder Colin/Type 
AT/Rothschild Bequest BM 1939-1/, and is similarly designated paralectotype. 

Salpinx oculata Moore was described from an unstated number of male specimens, 
from 'Philippines (Mindanao)', in the British Museum. The BMNH now possesses 
a single male which bears the following labels: /Type/Mindanao, 'Challenger' 83- 
62/Salpinx oculata <$ type Moore/BM type no. Rh 6768/. This specimen is hereby 
designated lectotype of Salpinx oculata Moore, and has been labelled accordingly. 



Doleschallia hexopthalmos hexopthalmos (Gmelin) [Nymphalidae] 

[Papilio polibete Cramer; Cramer, [1779] : 73, pi. 235, figs C, D. Misidentification.] 

Papilio Danai festivi sp., Zschach, 1788 : 90, no. 49*. 'Exoticus'. 

Papilio hexopthalmos Gmelin, 1790 : 2289, no. 890. LECTOTYPE $, 'extra Europam' [? 

Ambon] (NMI), here designated [examined]. 
Papilio hexopthalmos; O'Reilly, 1813 : 75, no. 49. 'India'. 
Papilio hexopthalmos Gmelin; Kirby, 1869 : 359. 
Doleschallia hexopthalmos (Gmelin); Kirby, 1871 : 194. 
Doleschallia amboinensis Staudinger, April 1885 : 104 [Feb. 1886], pi. 39 [Apr. 1885]. Holotype 

£, Ambon (MNHU). Synonymy by D'Abrera, 1971 : 228. 
Doleschallia crameri Distant, July 1885 : 41. Ambon (type-material lost). Synonymy by 

D'Abrera, 1971 : 228. 
Doleschallia hexopthalmos hexopthalmos (Gmelin); Fruhstorfer, 1912a : 561. 'Amboina'. 
Doleschallia hexopthalmos hexopthalmos (Gmelin); D'Abrera, 1971:228. 'Bum, Ambon, 

Serang, Saparua'. 

Papilio hexopthalmos Gmelin. A single female specimen in Dublin bears the 
labels /49/Hexopthalmos, Gmel [49] type specimen Doleschall [ia] Hexopthalmos, 
Gmel . . . [overlooked for 70 years, . . . [ujnfigured/, the latter being a badly cut 



BUTTERFLIES NAMED BY GMELIN 43 

label in Kirby's handwriting, on blue paper. This specimen, which is in moderate 
condition, is hereby designated lectotype of Papilio hexopthalmos, and has been 
labelled accordingly. 

Fruhstorfer (191 2 : 561) appears to have correctly interpreted Gmelin's species 
as an insect from Ambon; hexopthalmos is still regarded as a good species with a 
number of subspecies (D'Abrera, 1971 : 228). The classification of Doleschallia 
species is, however, perplexing (vide D'Abrera, op. cit.) ; anyone revising the group 
will have to pay particular attention to type-material, including that of hexopthalmos. 

Doleschallia crameri Distant. Distant (1885 : 41) recognized that Cramer had 
confused two species under the name polibete, describing first the male of one species 
(Cramer, 1779 : 71, pi. 234, figs D, E), and then a female of another (op. cit: 73, 
pi. 235, figs C. D). Distant restricted the 'type' of polibete to figs D, E on pi. 234, 
naming the insect illustrated on pi. 235, figs C, D, as crameri. In addition Distant 
referred to crameri material of both sexes collected on Ambon by Forbes. No 
crameri specimens attributable to either Cramer or Forbes are present in the Roths- 
child collection (BMNH), where they might have been expected. 

Papilio polibete Cramer. As noted above, Distant restricted the usage of this 
name to the species dealt with by Cramer (1779) on pages 71-72 and pi. 234, figs 
D, E. The material concerned evidently came from Ambon; the name polibete 
is currently in use for the subspecies of Doleschallia bisaltide (Cramer) recognized 
from Ambon, Ceram and Saparua (D'Abrera, 1971 : 266). No original specimens 
are to be found in the Rothschild collection, or apparently in Leiden (RNH) ; they 
are presumed lost or destroyed. In the absence of type-material of either crameri 
or polibete, Distant's usage should be followed; it may eventually be necessary to 
designate neotypes for one or both names. 



Zaretis itys (Cramer) [Charaxidae] 

Papilio itys Cramer, [1777] : 34, 149 (itus), pi. 119, figs F, G. LECTOTYPE $, Surinam 

(BMNH), here designated [examined]. 
Papilio isidora Cramer, [1779] : 72, pi. 235, figs A,B,E,F. i(J, 1 $ syntypes, Surinam (BMNH) 

[examined] . 
Papilio Danai festivi sp., Zschach, 1788 : 90, no. 55*. 'Exoticus'. 
Papilio strigosus Gmelin : 2290, no. 891. LECTOTYPE $, 'extra Europam' [? Surinam] 

(NMI), here designated [examined]. 
Papilio strigosus; O'Reilly, 1813 : 75, no. 55. 'India'. 
Siderone itylus Westwood, 1850 : 321. Holotype <3\ Brazil: Rio de Janeiro (BMNH) 

[examined]. Authorship and date: Hemming, 1941 : 456. 
Siderone zethus Westwood, 1850 : 321. Holotype £, Brazil: Para (BMNH) [examined]. 

Authorship and date: Hemming, 1941 : 463. 
Siderone ellops Menetries, 1855 : 88, pi. 3, fig. 1. g, $ syntypes, Nicaragua (ZI). 
Siderone strigosa (Gmelin); Kirby, 1869 : 359, 1871 : 280. 'Para'. 
Siderone isidora var. strigosa Staudinger, 1887 : 184, 185. Brazil: Rio de Janeiro to Rio 

Grande do Sul (MNHU). [Junior homonym of Papilio strigosus Gmelin, 1790.] 
Siderone isidora var. cacica Staudinger, 1887 : 184. Peru: Chanchamayo (MNHU). 
Zaretes isidora strigosa (Staudinger); Fruhstorfer, 1909 : 167. 'Sao Paulo to Sta. Catharina'. 
Z aretes itys pseuditys Fruhstorfer, 1909 : 166. 2 <$, 1 $ syntypes, Brazil: Espirito Santo 

(BMNH) [examined]. 



44 R- I- VANE-WRIGHT 

Zaretes isidora anzuletta Fruhstorfer, 1909 : 167. 1 <$, 1 Q syntypes, Mexico (BMNH) 

[examined] . 
Zaretes isidora russeus Fruhstorfer, 1909 : 167. 1 $ syntype, Colombia (BMNH) [examined]. 
Zaretes isidora vulpecula, Fruhstorfer, 1909 : 167. 3 o*, 1 $ syntypes, Brazil: Bahia (BMNH) 

[examined] . 
Zaretes isidora leopoldina Fruhstorfer, 1909 : 167. 1 $ syntype, Brazil: Espirito Santo 

(BMNH) [examined]. 
Zaretes isidora vulpina Fruhstorfer, 1909 : 167. 7 ^, 1 $ syntypes, Paraguay (BMNH) 

[examined] . 
Siderone strigosus (Gmelin) ; Weeks, 191 1 : xii. 'Venezuela: Suapure'. 
Zaretes isidora zethus (Westwood) ; Rober, 1916 : 578. ('= strigosus Gmelin?; lower Amazon 

and Paraquay'). 
Zaretes isidora strigosus (Gmelin); Talbot, 1928 : 209. 'Mato Grosso'. 
Zaretis itys strigosus (Gmelin); Stichel, 1939 : 726. 'Para'. 

Zaretis itys strigosa (Staudinger) ; Stichel, 1939 : 726. (= strigosus Gmelin?; Rio de Janeiro'). 
Anaea (Zaretis) itys (Cramer); Comstock, 1961 :3o. (All available names carried in synonymy). 

Papilio strigosus Gmelin. A single female specimen in Dublin bears the label 
/55 Strigosus, Gmel. type specimen/, in Kirby's handwriting, on blue paper. This 
specimen is hereby designated lectotype of Papilio strigosus Gmelin, and has been 
labelled accordingly. It is in rather poor condition, badly worn and variously 
repaired; one 'antenna' consists of a piece of bristle. As discussed below, this 
specimen most probably came from Surinam. 

Comstock (1961), in his treatment of Anaea and allies followed, according to 
many workers, a very conservative line, demoting Zaretis to subgeneric rank. 
Further, while admitting evidence of subspeciation in Zaretis itys, he treated all 
available names applied to this species as synonymous, stating that 'with the present 
confused state of the literature and the unsatisfactory working material no attempt 
is made at the present time to assign the names properly' (Comstock, 1961 : 31). 

Having examined the literature as pertaining just to the name strigosus Gmelin, 
I feel sympathy for Comstock's view. As can be seen from the synonymy presented 
above, the name strigosa or strigosus has been variously applied to material from 
Venezuela, Para, Mato Grosso and southern Brazil. Most confusingly, Staudinger 
(1887) named strigosa from South Brazil, evidently unsure whether his strigosa 
was the same as strigosus Gmelin or not. Clearly in this case strigosa Staudinger 
should be treated as an invalid homonym; the oldest valid name for such a sub- 
species, in any case, is itylus Westwood. This leaves strigosus Gmelin as a name 
most commonly applied to material of itys from the lower Amazon (Para) . Having 
examined the lectotype, however, I see no reason why the name should not be 
applied to the Guyana/Surinam region, the most likely locality for the Leske material. 
The lectotype is perhaps closest in appearance to material of itys from Trinidad 
(in NMI and BMNH). 

I refrain at this time from designating further lectotypes for the names pertaining 
to itys, as listed above in synonymy, with the exception of itys itself. It seems likely 
that Comstock's conservative approach will be superceded; Rydon (1971) has 
recently reinstated Zaretis to full generic rank. Those who may wish to rework 
the infraspecific and infrasubspecific taxonomy of itys will find further references 
in Stichel (1939) and Comstock (1961). In the course of any thorough revision, 



BUTTERFLIES NAMED BY GMELIN 45 

the lectotype of strigosus Gmelin will have to be re-examined. If possible, however, 
I suspect that the best course will be to apply strigosus to a Surinam insect, thereby 
removing this confusing name from the classification by placing it in the strict 
synonymy of itys Cramer (assuming that only one species is involved in this complex). 
A lectotype designation for itys, the senior name, is appended. 

Papilio itys Cramer was described from an unstated number of female specimens 
from Surinam. A single female in the Rothschild collection (BMNH) which bears 
the labels /Surinam Coll Lenep/Felder Colin/ compares closely with Cramer's original 
illustrations. This specimen is hereby designated lectotype of Papilio itys Cramer, 
and has been labelled accordingly (type no. Rh 17307). 



Hypolimnas pandarus pandarus (Linnaeus) [Nymphalidae] 

Papilio pandarus Linnaeus, 1758 1461 ; Clerck, 1764 : pi. 26, fig. 2. LECTOTYPE $, 'in Indus' 

[Ambon] (MLII), here designated. Type-locality: see Corbet, 1941 : 15; Corbet, 1949 : 193, 

197. 
Papilio pipleis Linnaeus, 1758 1476; Clerck, 1764 :pl. 26, fig. 2. LECTOTYPE $, 'in Indiis' 

[Ambon] (MLU), here designated. Synonymy from Kirby, 1871 : 225. Type-locality: 

see Corbet, 1941 : 14; Corbet, 1949 : 193, 197. 
Papilio calisto Cramer, 1775 : 37, pi. 24, figs A, B. LECTOTYPE <J, 'Africa' [? Ambon] 

(BMNH), here designated [examined]. Synonymy from Kirby, 1871 : 225. 
Papilio pipleis Linnaeus; Cramer, 1775 : 93, pi. 60, figs A, B. ['Amboina'; compared with 

calisto.] 
Papilio Nymphales Gemmati sp., Zschach, 1788 : 90, no. 58b*. 'Exoticus'. 
Papilio lacteolus Gmelin, 1790 : 2290, no. 892. LECTOTYPE §, 'extra Europam' [? Ambon] 

(NMI), here designated [examined]. Synonymy from Kirby, 1869 : 359. 
Papilio lacteolus; O'Reilly, 181 3 : 75, no. 58b. 'India'. 
Diadema pandarus (Linnaeus); Kirby, 1869 : 359. 

Hypolimnas pandarus (Linnaeus); Kirby, 1871 : 225. 'Amboina, Ceram'. 
Hypolimnas pandarus pandarus (Linnaeus); Fruhstorfer, 1912a : 554. 'Amboina, Saparua, 

Ceram'. 
Hypolimnas pandarus pandarus (Linnaeus); D'Abrera, 1971 : 224. 
[Hypolimnas hewitsoni (Wallace); D'Abrera, 1971 : 224. Misidentification.] 

Papilio lacteolus Gmelin. A single female in Dublin bears the labels /58b Pan- 
darus. type of Lacteolus, GmeL/21. 5. 6. 79./, the former in Kirby's handwriting. 
This specimen is hereby designated lectotype of Papilio lacteolus Gmelin, and has 
been labelled accordingly. In quite good condition, the lectotype is very similar 
to the specimen erroneously illustrated by D'Abrera (1971 : 225) as the female 
of 'Hypolimnas pandarus hewitsoni', from 'Kai'. The latter is in fact a pandarus 
female from Ceram; an exactly similar form, however, occurs on Ambon, which 
seems the most likely type-locality for lacteolus. True hewitsoni (Wallace) is a 
subspecies of H. deois (Hewitson). 

Corbet (1941; 1949) considered the Linnaean species pandarus and pipleis to 
be from Ambon. Corbet further considered both these names to be based on the 
same type-specimen, illustrated by Clerck (1764), and thus objectively synonymous. 
For completeness I append lectotype designations for these, to finally establish 
this synonymy, and for calisto Cramer, the remaining synonym. 



46 R. I. VANE-WRIGHT 

Papilio pandarus Linnaeus. Corbet (1949 : 197) summarizes his work on the 
type-material of this species. I hereby designate the specimen in MLU, as dealt 
with by Corbet (loc. cit.) and illustrated by Clerck (1764 : pi. 26, fig. 2, as Papilio 
pipleis), lectotype of Papilio pandarus Linnaeus. 

Papilio pipleis Linnaeus. Corbet (1949 : 197) summarizes his work on the 
type-material of this species. I hereby designate the specimen in MLU, as dealt 
with by Corbet (loc. cit.) and illustrated by Clerck (1764 : pi. 26, fig. 2), lectotype 
of Papilio pipleis Linnaeus, which is thereby finally established as an objective 
synonym of pandarus L. 

Papilio calisto Cramer was described from an unstated number of specimens 
from 'Africa', in the collection of W. van der Meulen. The BMNH possesses a 
single male received in the Rothschild collection which corresponds well with 
Cramer's original illustration, and bears the following labels: /No. 40. calisto. 
Cr.i.24.A.B./Felder Colin/Rothschild Bequest BM 1939-1/. The former label 
is in the characteristic style believed to be associated with Cramer material. This 
specimen is hereby designated lectotype of Papilio calisto Cramer, and has been 
labelled accordingly (type no. Rh 17300). 

Antirrhea bifasciatus (Gmelin) [Satyridae ) 

Papilio Nymphales Gemmati sp., Zschach, 1788 : 91, no. 59*. 'Exoticus.' 

Papilio bifasciatus Gmelin, 1790 : 2290, no. 893. 'Extra Europam' [? Surinam] (formerly 

in NMI, type-material destroyed). 
Papilio bifasciatus; O'Reilly, 1813 : 75, no. 59. 'India'. 
Papilio bifasciatus Gmelin; Kirby, 1869 : 359. 
Antirrhaea bifasciatus (Gmelin); Kirby, 1871 : 642; Weymer, 1910 : 182 (footnote). 

Papilio bifasciatus Gmelin. Kirby (1869 : 360) states of this species 'One of 
the Satyriqae. A fragment of one of the types is still in existence; but I have not 
yet succeeded in identifying it'. Two years later (Kirby, 1871 : 642) he placed 
bifasciatus in Antirrhea, to follow the well known species taygetina Butler. My 
searches in Dublin for the 'fragment' of the type proved fruitless; for the reasons 
discussed in detail in the Historical Review above (p. 24), I prefer to retain this 
name as a species inquirenda of the genus Antirrhea. 

Papilio sulpitia Gmelin [Pfamily] 

Papilio Nymphales Phalerati sp., Zschach, 1788 : 92, no. 88*. 'Exoticus'. 

Papilio sulpitia Gmelin, 1790 : 2336, no. 894. 'Extra Europam' (formerly in NMI, type-material 

destroyed). [Junior homonym of Papilio sulpitia Cramer, 1779, and Papilio sulpitia Stoll, 

1780.] 
Papilio sulpitia; O'Reilly, 1813 : 76. 'India'. 
Papilio sulpitia Gmelin; Kirby, 1869 : 360; 1871 : 637; Sherborn, 1902 : 948. 

Papilio sulpitia Gmelin. The type-material of this species was evidently des- 
troyed sometime between 1813 and 1869. Kirby (1869 : 360) repeats the original 
Latin description; probably unrecognizable, this forgotten name is in any case 
invalid, being a junior primary homonym of both Papilio sulpitia Cramer and 



BUTTERFLIES NAMED BY GMELIN 47 

P. sulpitia Stoll. Both the latter were, however, described within 'Papilio Nym- 
phales Phalerati\ as was sulpitia Gmelin; the original description of Zschach (con- 
siderably shortened and modified by Gmelin) could apply to Cramer's species, 
the Chinese Ladoga sulpitia, but not to Stoll's Metamorpha elissa Hubner (= sulpitia 
Stoll), from South America. Conceivably Gmelin intended to refer to Cramer's 
original sulpitia, having perhaps identified Zschach's Papilio no. 88 with it. How- 
ever, as Gmelin did not make any such reference, the name sulpitia Gmelin should 
be treated as suggested above. 



Papilio vidua Gmelin [Pfamily] 

Papilio Nymphales Phalerati sp., Zschach, 1788 : 92, no. 89*. 'Exoticus'. 

Papilio vidua Gmelin, 1790 : 2336, no. 895. 'Extra Europam' (formerly in NMI, type-material 

destroyed). [Junior homonym of Papilio vidua Miiller, 1764.] 
Papilio vidua; O'Reilly, 1813:76, no. 89. 'India'. 
Papilio vidua Gmelin; Kirby, 1869 : 360; 1871 : 637; Sherborn, 1902 : 1044. 

Papilio vidua Gmelin. The type-material was evidently destroyed or lost 
between 1813 and 1869. Kirby (1869 : 360) repeats the original description; 
probably unrecognizable, this forgotten name is invalid, being a junior primary 
homonym of Papilio vidua Miiller. In this case there is no question of Gmelin 
having failed to give a reference to the original name; vidua Miiller and vidua Gmelin 
are unquestionably different, Miiller's species being referable to the European 
Satyrid Aphantopus hyperantus (Linnaeus) . 



Thecla cupentus (Stoll) s. str. [Lycaenidae] 

Papilio cupentus Stoll, [1781] : 93, pi. 337, figs F, G. LECTOTYPE <J, Surinam (BMNH), 
here designated [examined]. Correct date and authorship: Hemming, 1958 :43; Brown, 
1941. 

Papilio Plebeii Rurales sp., Zschach, 1788 : 93, no. 123*. 'Exoticus.' 

Papilio annulatus Gmelin, 1790 : 2359, no. 896. LECTOTYPE $, 'extra Europam' [? 
Surinam] (NMI), here designated [examined]. Synonymy by Draudt, 1920 : 769. 

Papilio annulatus; O'Reilly, 1813 : 76, no. 123. 'Surinam'. 

Papilio annulatus Gmelin; Kirby, 1869 : 361. 

Thecla annulatus (Gmelin); Kirby, 1871 : 399. 

Thecla cupentus (Cramer); Draudt, 1920 : 769; Comstock & Huntington, 1959 : 72. 

Papilio annulatus Gmelin. A single male specimen in Dublin bears the labels 
/123/123 Annulatus, Gmel. type/, in Kirby's handwriting; this specimen is hereby 
designated lectotype of Papilio annulatus Gmelin, and has been labelled accordingly. 
The specimen is in rather poor condition, lacking antennae and large portions 
of the hind wings; the abdomen has now been dissected (preserved on separate 
75 mm x 25 mm glass slide). The lectotype is evidently from the Surinam region; 
it is amusing to note that while Zschach and Gmelin gave their usual 'Exoticus'' 
and 'extra Europam' respectively, O'Reilly forsook his usual 'India' for 'Surinam', 
this change, however, probably being accidental. 

I am indebted to Dr H. K. Clench for much help and information concerning 



48 R. I. VANE-WRIGHT 

this species. He has pointed out to me that cupentus must eventually be removed 
from the omnibus 'Thecla\ probably requiring the erection of a new genus to receive 
it. Further, he notes that the appearance of cupentus from the Guyana region is 
rather different compared with material from other areas in South America, perhaps 
indicating that this species will eventually be broken up into subspecies or siblings. 
To check this point Dr Clench sent me drawings of the male genitalia of a supposed 
cupentus from Colombia. I have compared this with dissections of the lectotypes 
of both annulatus and cupentus, finding all three to be in no way significantly differ- 
ent. I thus regard the latter two names as strictly synonymous, pertaining to 
material from the Guyana region [cupentus Stoll s. str.) ; the Colombian specimen 
would appear to represent, at most, only a separate subspecies. The lectotype 
designation for cupentus follows. 

Papilio cupentus Stoll was described from an unstated number of male specimens 
from 'Suriname', apparently lent to Stoll by W. van der Meulen. A single male 
specimen in the BMNH now bears the following labels: /Surinam. Lenip/Felder 
Colin/Rothschild Bequest BM 1939-1/. As explained elsewhere, a number of 
apparent Cramer and Stoll specimens reached the Rothschild collection via van 
Lennep and Felder. I have compared this specimen with Stoll's published work, 
and the original paintings, with which it compares quite closely, allowing for some 
'artistic licence' (as with all Cramer and Stoll illustrations, they are somewhat 
stylized). This specimen is hereby designated lectotype of Papilio cupentus Stoll, 
and has been labelled accordingly (type no. Rh 17301; slide no. Rh 11176). 

Cupido oculatus (Gmelin) [Lycaenidae] 

Papilio Plebeii Rurales sp., Zschach, 1788 : 93, no. 131*. 'Eur'. 

Papilio oculatus Gmelin, 1790 : 2359, no. 897. 'In Europa' (formerly in NMI; type-material 

destroyed) . 
Papilio oculatus; O'Reilly, 1813 : 77, no. 131. 'Europe'. 
Papilio oculatus Gmelin; Kirby, 1869 : 361. 
Cupido oculatus (Gmelin); Kirby, 1871 : 375. 

Papilio oculatus Gmelin. The type-material of this species was evidently lost 
or destroyed between 1813 and 1869. Kirby (1869 : 361) repeats the original 
description and indicates (under lunulatus Gmelin) that this species is a European 
Lycaenid, probably belonging either to the anon-group (Maculinea van Eecke) or 
Polyommatus Latreille. Two years later Kirby (1871 : 375) placed oculatus in 
his omnibus treatment of Cupido Schrank (to include Polyommatus), near avion 
(L.). The name oculatus seems subsequently to have been unused. It has never 
been recognised as pertaining to any known species. Despite Kirby's claim that 
it may prove possible to identify, no one has been able to do so yet. 

Cupido lunulatus (Gmelin) [Lycaenidae] 

Papilio Plebeii Rurales sp., Zschach, 1788 : 93, no. 132*. 'Eur'. 

Papilio lunulatus Gmelin, 1790 : 2359, no. 898. 'In Europa' (formerly in NMI; type-material 
destroyed). 



BUTTERFLIES NAMED BY GMELIN 49 

Papilio lunulatus; O'Reilly, 1813 : 76, 77, nos 130, 132. 'Europe'. 
Papilio lunulatus Gmelin; Kirby, 1869 : 361. 
Cupido lunulatus (Gmelin); Kirby, 187 1 : 375. 

Papilio lunulatus Gmelin. The type-material has evidently been destroyed. 
All comments applied to Papilio oculatus above apply equally to this species, but 
see also entry below (p. 54) for Papilio Plebeii Rurales sp., Zschach no. 130*. 

Jemadia hospita hephaestos (Plotz) stat. n., nom. rev. [Hesperiidae] 

(PI. 6, fig. 24) 

Papilio Plebeii Urbicolae sp., Zschach, 1788 : 94, no. 154*. 'Exoticus'. 

Papilio aethiops Gmelin, 1790 : 2360, no. 899. LECTOTYPE^, 'extra Europam' [? Surinam] 
(NMI), here designated [examined). Incorrectly synonymized with Jemadia menechmus 
(Mabille) by Evans, 1951 : 52. [Junior homonym of Papilio aethiops Esper, 1779.] 
Syn. n. 

Papilio aethiops; O'Reilly, 1813 : 77, no. 154. 'Africa'. 

Papilio aethiops Gmelin; Kirby, 1869 : 361. 

? Pyrrhopyge aethiops (Gmelin); Kirby, 1871 : 587. 

Pyrrliopyga hephaestos Plotz (Moschler in litt.), 1879 1521. Holotype 9. Surinam [; Para- 
maribo] (MNHU, Berlin) [examined]. 

Pyrrhopyge hephaestus Moschler, 1883 : 324. Holotype 9. Surinam: Paramaribo (MNHU) 
[examined]. Based on the same type-specimen, the name hephaestus Moschler is an objective 
synonym of hephaestos Plotz. 

? Pyrrhopyge aethiops (Gmelin); Mabille, 191 2 : 7. 

[Jemadia hospita ulixes (Plotz); Evans, 1915 : 51. Misidentification.] 

Papilio aethiops Gmelin; Evans, 1951 : 52 (primary homonym of aethiops Esper). [Incorrectly 
placed as a synonym of Jemadia menechmus (Mabille).] 

Papilio aethiops Gmelin. A single male specimen in Dublin bears the label 
/154 Aethiops, Gmel. Type specimen/, in Kirby's handwriting; this specimen is 
hereby designated lectotype of Papilio aethiops Gmelin, and has been labelled 
accordingly. It is in rather poor condition. The abdomen has been dissected 
and is now preserved on a separate 75 mm x 25 mm glass slide. The type-locality 
of aethiops Gmelin is almost certainly Surinam (O'Reilly was misled by the name). 
However, although the few specimens in the BMNH which correspond to aethiops, 
as dealt with by Evans (1951 : 51) under Jemadia hospita ulixes, mostly bear the 
designation 'Surinam', all are very old, 18th century, as is the type of aethiops. 
The name aethiops Gmelin is an invalid homonym; the only suitable replacement 
that I have been able to find is hephaestos Plotz. However, it is with some doubt 
that I here refer the 18th century aethiops Gmelin material to the latter name, 
which is based on a 19th century specimen (PI. 6, fig. 24) and does not correspond 
precisely with true aethiops, or any other material that I have seen. The type of 
hephaestos is, unfortunately, a female; it may eventually prove necessary, or the 
better course, to propose a new name for aethiops Gmelin. The species requires 
the acquisition of fresh material and careful re-examination. 

The following notes on the synonymies of Jemadia menechmus, J. hospita and 
/. hewitsonii stem from examining Evans' (1951) work with respect to the synonymy 
of aethiops Gmelin. 



5 o R. I. VANE-WRIGHT 

Jemadia menechmus (Mabille) [Hesperiidae] 

Evans (1951 : 52) identified aethiops Gmelin with this name. As shown above, 
aethiops pertains to /. hospita (Butler) , and should be removed from the synonymy 
of menechmus. 

Jemadia hospita hospita (Butler) [Hesperiidae] 
(PI. 6, fig. 23) 

Pyrrhopyga hospita Butler, 1877 : 128. LECTOTYPE <J, Peru: Ucayali (BMNH), here 

designated [examined]. 
Pyrrhopyga ulixes Plotz, 1879 : 521. LECTOTYPE^, 'Suriname' [? true locality] (MNHU), 

here designated [examined]. Syn. n. 

Evans (1951 : 51) dealt with material attributable to true aethiops Gmelin as 
subspecies /. hospita ulixes (Plotz). However, examination of the ulixes type 
(PI. 6, fig. 23) shows it to be very similar to hospita Butler; it seems possible that 
it came from the upper Amazon region rather than Surinam as originally indicated. 

Pyrrhopyga hospita Butler was described from 'several examples' collected at 
Ucayali, Peru, by Walter Davis. A single male specimen now remains in the 
BMNH and bears the labels /Type H.48/Peru. W. Davis. 77-52/P. hospita Butler 
Type/; this specimen is hereby designated lectotype of Pyrrhopyga hospita Butler 
and has been labelled accordingly (type no. Rh 17306). 

Pyrrhopyga ulixes Plotz was described from an unstated number of specimens 
from 'Surinam'. A single male specimen in the MNHU bears the labels /Type/ 
Surinam VoliKman/Ulixes Plotz*/Ulixes Plotz Stett.Ent.Zeit. 1879, p.521/4931/ 
9 : 16/, and is hereby designated lectotype of Pyrrhopyga ulixes Plotz (PI. 6, fig. 
23) ; it has been labelled accordingly. 

Jemadia hewitsonii brevipennis Schaus [Hesperiidae] 

Jemadia brevipennis Schaus, 1902 : 425. Holotype, Brazil: Sao Paulo (USNM), type no. 

5962. 
? Jemadia lisetta Mabille & Boullet, 1908 : 195, 197. 1 $ syntype, Peru (MNHN), [examined]. 

Incorrectly synonymized with /. hospita ulixes by Evans, 1951 : 51. 
Jemadia hewitsonii brevipennis Schaus; Evans, 1951 : 54. 'Bolivia, Mato Grosso, Sao Paulo'. 

As noted above, Evans (1951 : 51) dealt with true aethiops Gmelin as /. hospita 
ulixes, to which he also attributed lisetta Mabille & Boullet. The latter was described 
from three males in the Boullet collection from Peru and another male in the Mabille 
collection, from Bolivia. Evans effectively claimed to have identified the latter 
specimen in the BMNH collections, noting that it was labelled 'lisetta' by Mabille, 
had no locality data, and was probably from Surinam. This specimen he regarded 
as the 'type', and it formed the basis of his synonymy. However, Mabille named 
a considerable amount of material in the BMNH collections, often inaccurately. 
I do not consider there to be sufficient evidence to regard this specimen as the 
'Bolivian' example noted in the original description. I have examined one of the 



BUTTERFLIES NAMED BY GMELIN 51 

syntypic Peru males (PL 6, fig. 25) of the Boullet collection, now in the MNHN, 
and find it to be extremely similar to /. hewitsonii brevipennis Schaus, as dealt 
with by Evans (1951 : 54). This, however, is rather surprising, since one might 
expect hewitsonii material from Peru to fit the nominotypical subspecies. A 
cursory examination of hewitsonii material as dealt with by Evans suggests that 
it is possible that two species, with overlapping ranges may be involved; the whole 
hewitsonii complex needs careful re-examination, as indeed, does the whole species 
taxonomy of Jemadia, Evans' account seemingly being inadequate in a number of 
ways. 

Autochton bipunctatus (Gmelin) [Hesperiidae] 

(PL 6, fig. 26) 

Papilio Plebeii Urbicolae sp., Zschach, 1788 : 94, no. 160*. 'Exoticus'. 

Papilio bipunctatus Gmelin, 1790 : 2360, no. 900. NEOTYPE <§, French Guiana (BMNH), 

here designated [examined]. 
Papilio bipunctatus; O'Reilly, 1813 : 77, no. 160. 'India'. 
Papilio bipunctatus Gmelin; Kirby, 1869 : 362. 

Thymele bipunctatus (Gmelin); Kirby, 1871 : 572. Cecrops neis Geyer given as a synonym. 
Cecropterus bipunctatus (Gmelin); Moschler, 1877 : 344; Kirby, 1880 : 329. 
Cecropterus neis (Geyer); Plotz, 1882 : 261. 'Laguayra'; bipunctatus given as a synonym of 

nets. 
Cecropterus orontes Plotz, 1882 : 261. Venezuela: La Guaira (type-material lost). Synonymy 

by Evans, 1952 : 125. 
Cecropterus bipunctatus (Gmelin); Mabille, 1883 : lv. As separate species from neis. 
Cecropterus zonilis Mabille, 1883 : lvi. LECTOTYPE <J, Colombia (BMNH), here designated 

[examined]. Synonymy from Evans, 1952 : 125. 
Cecropterus bipunctatus (Gmelin); Watson, 1893 : 32. 
Cecropterus neis (Geyer); Godman & Salvin, 1894 : 328. C. bipunctatus (Gmelin) given as a 

doubtful synonym. 
Cecropterus bipunctatus (Gmelin); Mabille, 1903 : 29; Kaye, 1904 : 211. 
Cecropterus bipunctatus (Gmelin) ; Draudt, 1922 : 870. ? Misidentihcation according to Evans, 

1952 : 125. 
Autochton bipunctatus (Gmelin); Evans, 1952 : 125. 
Cecropterus bipunctatus (Gmelin); Barcant, 1970 : 298. 

Papilio bipunctatus Gmelin. The specimen of bipunctatus studied by Kirby 
(1869 : 362; 1871 : 572) is no longer to be found in Dublin. Kirby (1880 : 329) 
lists one specimen of this species, but as usual this does not refer to type-material 
(see discussion of Papilio argyrios, p. 29). Assuming Kirby (1871) to be essentially 
correct in his treatment of this species, bipunctatus Gmelin may be regarded as a 
species of the group now dealt with as Autochton Hiibner; the most likely type- 
locality for bipunctatus would have been Surinam. 

Kirby (1871) gave bipunctatus as a senior synonym of Cecrops neis Geyer. Mabille 
appears to have been the first to use both of these names for supposedly distinct 
species. Draudt (1922), according to Evans (1952), 'misidentified' bipunctatus, 
probably dealing with the species longipennis Plotz under that name. Very probably 
many of the authors dealing with bipunctatus as a separate species have variously 
misidentified it in Evans' sense. Evans (1952), while dealing with neis and longi- 



52 R. I. VANE-WRIGHT 

pennis as separate species, treats bipunctatus as the senior synonym for orontes 
Plotz and zonilis Mabille. Barcant (1970) lists bipunctatus in the sense of Kaye 
(1904). 

The need for a primary type-specimen for Papilio bipunctatus Gmelin is apparent. 
To satisfy this requirement in a manner which least upsets existing usage, one of 
the male specimens studied by Evans (1952 : 125) from French Guiana, is here 
designated neotype of Papilio bipunctatus Gmelin. This specimen, in the BMNH, 
bears the labels /Guyane Francaise Collection C. Bar/R.Oberthiir Coll. BM 1931-136/ 
Papilio bipunctatus Gmelin, Neotype $ det. R.I.V.-W. 1974./. This neotype is 
illustrated on PI. 6, fig. 26. 

Cecropterus orontes Plotz. I have been unable to locate any type-material of 
this name. Apparently no type-specimens exist in Berlin, where they might have 
been expected; possibly the original material is destroyed. 

Cecropterus zonilis Mabile was described from an unstated number of specimens 
of unstated sex, from 'Columbia'. A single male now in the BMNH bears the 
labels /Colombia/zonilis Mb. orontes PI. /type/Ex Oberthur Coll. BM 1927-3/. 
This specimen, referred to by Evans (1952 : 125) as the 'type', is hereby designated 
lectotype of Cecropterus zonilis Mabille, and has been labelled accordingly (type 
no. Rh 17302). A further pair of specimens, possibly part of the original series, 
are not designated paralectotypes at this time. 

Ablepsis fenestratus (Gmelin) [Hesperiidae] 
(PL 6, fig. 27) 

Papilio Plebeii Urbicolae sp., Zschach, 1788 : 95, no. 161*. 'Exoticus'. 

Papilio fenestratus Gmelin, 1790 12360, no. 901. NEOTYPE^, French Guiana: Nouveau 

Chantier (MNHN), here designated [examined]. 
Papilio fenestratus ; O'Reilly, 1813 : 77, no. 161. 'India'. 
Papilio fenestratus Gmelin; Kirby, 1869 : 362. 
Plesioneura fenestratus (Gmelin); Kirby, 1871 : 621. 
Telemiades acutipennis Mabille & Boullet, 1912 : 120. Holotype <3\ French Guiana: Nouveau 

Chantier (MNHN) [examined]. [Synonymy originally from Evans, 1952 : 162; becomes 

objective synonym of fenestratus as result of action taken here.] 
Papilio fenestratus Gmelin; Evans, 1949 : 477. 
Ablepsis fenestratus (Gmelin); Evans, 1952 : 162. 

Papilio fenestratus Gmelin. The original type-material has been destroyed. 
Kirby (1869 : 362) states that the type was then still in existence, but so broken 
that there was little chance of identification. Two years later, however, he placed 
it in Plesioneura (Kirby, 1871 : 621). Evans (1949 : 477) was unable to fit the 
original description of fenestratus to any true Plesioneura known, or any of the 
other Old World species then under review, but noted it to be in accord with the 
S. American Telemiades acutipennis Mabille & Boullet. Evans also noted that 
the type of fenestratus was no longer in existence, which I am able to confirm after 
studying the collection at Dublin. Evans (1952 : 162) finally placed fenestratus 
in Ablepsis, with acutipennis as a synonym. Evans studied only a single male, 
in the BMNH from French Guiana, and which I have closely compared with the 



BUTTERFLIES NAMED BY GMELIN 53 

holotype of acutipennis (PI. 6, fig. 27). I can confirm Evans' identification, and 
to stabilise existing usage, I here designate the holotype of Telemiades acutipennis 
Mabille & Boullet as additionally the neotype of Papilio fenestratus Gmelin. This 
male specimen, now in the MNHN, bears the data label /French Guiana, Nouveau 
Chantier, E. le Loult/, and is illustrated on PI. 6, fig. 27. 



REPLACEMENT NAMES PROPOSED BY GMELIN 

Syntarucus pirithous (Linnaeus) [Lycaenidae] 

Papilio pirithous Linnaeus, 1767 : 790, no. 235. Algeria (type-material lost?). 

Papilio philiasus Linnaeus, 1767 : 790, no. 233. Algeria (type-material lost?). Synonymy 

by Kirby, 1871 : 351. 
Papilio telicanus Lang, 1789 : 47. France: southern (depository of type-material unknown). 

Synonymy by Higgins & Riley, 1970 : 251. 
Papilio barbarus Gmelin, 1790 : 2352, no. 235. [Proposed as a replacement name for Papilio 

pirithous Linnaeus, 1767.] 

Papilio barbarus Gmelin. As already shown (p. 25), Gmelin's number 235 was a 
reference to that of Linnaeus, and this name is an invalid replacement for pirithous 
Linnaeus, 1767. 

Kirby (1870 : 149, 150) tentatively synonymized P. pirithous and P. philiasus 
as the male and female of the same species, adding that the latter might also be 
conspecific with P. amyntas Fabricius. A year later Kirby (1871 : 351) confirmed 
the synonymy of P. philiasus, P. pirithous and P. barbaras and acted as first 
reviser in giving priority to the first over the second. At the same time he moved 
P. amyntas Fabricius to the synonymy of another species (p. 356). Finally, citing 
the same reference, he placed it as a Hesperiid (p. 605). By a coincidence, the 
true P. amyntas Fabricius (also a junior homonym) features under the next species 
below. Strong support for Kirby's synonymy of P. philiasus and P. pirithous 
as sexual dimorphs can be drawn from their original descriptions, in which for 
both Linnaeus wrote 'Habitat Algiriae, Brunniche'. Martin T. Briinnich (1737- 
1827) of Copenhagen was a correspondent of Linnaeus. He probably 'communica- 
ted' the descriptions, and his insects were probably captured flying together. 

As far as can be judged at present, Syntarucus pirithous (Linnaeus) has as subjec- 
tive synonyms Papilio philiasus Linnaeus and Papilio telicanus Lang, with Papilio 
barbarus Gmelin an objective synonym. However, no type-material has been 
traced, and further research is desirable. Staudinger & Rebel (1901 : 76, no. 
530 and n) provisionally linked the first three names. 

Polygonus leo leo (Gmelin) [Hesperiidae] 

Papilio amyntas Fabricius, 1775 : 533, no. 384. 'In America' (2 specimens in Kiel; see Zimsen, 

1964 : 518). [Homonym of Papilio amyntas Poda, 1761.] 
Papilio leo Gmelin, 1790 : 2363, no. 836. [Proposed as replacement name for Papilio amyntas 

Fabricius, 1775.] 
Polygonus leo leo (Gmelin); Evans, 1952 : 54. 



54 R. I. VANE-WRIGHT 

Gmelin evidently noted the homonymy of Papilio amyntas Fabricius, and pro- 
posed leo as a new name for it. The reader is referred to Evans (1952 : 54) for the 
full synonymy of this neotropical skipper. Gmelin also noted the name Papilio 
phocus Cramer, 1777, as a possible synonym. P. phocus is now regarded as a 
valid species, being the type-species of the genus Nascus Watson. 

Perichares philetes philetes (Gmelin) [Hesperiidae] 

Papilio covidon Fabricius, 1775 : 533, no. 385. 'In Iamaica' [Jamaica] (2 specimens in Glasgow; 

see Zimsen, 1964 : 518). [Homonym of Papilio coridon Poda, 1761.] 
Papilio philetes Gmelin, 1790 : 2364, no. 842. [Proposed as replacement name for Papilio 

coridon Fabricius, 1775.] 
Perichares philetes philetes (Gmelin); Evans, 1955 : 255. 

Gmelin evidently noted the homonymy of Papilio coridon Fabricius, and put 
forward the name philetes as a replacement. Gmelin also included the name Papilio 
talus Cramer, 1777, with this species, but talus is now regarded as a valid species 
of the genus Astraptes Hiibner. Papilio coridon Fabricius is the type-species of 
the genus Perichares Scudder. Hemming (1967 : 352) states that philetes Gmelin 
is the oldest valid name 'subjectively applicable to the type-species of this genus'. 
In my opinion, philetes was proposed as a replacement name, and is therefore objec- 
tively applicable to the type-species of Perichares. 



BUTTERFLIES DESCRIBED IN MUSEUM LESKEANUM BUT NOT NAMED 

BY GMELIN AS NEW SPECIES 

? Jamides celeno (Cramer) [Lycaenidae] 

Papilio Plebeii Rurales sp., Zschach, 1788 : 93, no. 122 (? celerio [sic]). 'Exoticus'. 
? Papilio celeno Cramer; Gmelin, 1790 : 2339, no. 705. 'Surinamo'. 
Papilio celerio [sic]; O'Reilly, 1813 : 76, no. 122. 'Surinam'. 

Papilio sp. no. 122, Museum Leskeanum. Zschach, in his description, cryptically 
indicates that this species may be Papilio 'celerio' (after Fabricius), correctly celeno 
Cramer (1775 : 51, pi. 31, figs C, D), described as from Surinam. Cramer's species 
is now recognized as a widespread member of the Indo-Australian genus Jamides. 
Gmelin (1790 : 2339) repeats Zschach's doubtful identity when dealing with celeno. 
O'Reilly (1813 : 76) places the species without question as 'celerio', thus repeating 
the incorrect spelling of Zschach, which stemmed originally from that of Fabricius. 

Assuming that the identity with the rather distinctive celeno is essentially correct, 
it would seem very likely that Papilio no. 122 actually referred to Jamides celeno 
sandy a Fruhstorfer, the subspecies to be found on Ambon. The original material 
has apparently been destroyed,. 



? 



Cupido lunulatus (Gmelin) [Lycaenidae] 



Papilio Plebeii Rurales sp., Zschach, 1788 : 93, no. 130*. 'Eur[ope]\ 
Papilio lunulatus; O'Reilly, 1813 : 76, no. 130. 'Europe'. 



BUTTERFLIES NAMED BY GMELIN 55 

Papilio sp. no. 130*, Museum Leskeanum. Gmelin apparently did not take this 
species into account. O'Reilly referred it to Papilio lunulatus, named by Gmelin 
on the basis of Papilio no. 132 of Zschach. As dealt with above, lunulatus itself 
has never been recognized, and although the specimens of Papilios 130 and 132 
were evidently both in Dublin in 1813, neither has survived to the present time. 



ZSCHACH, CRAMER, AMBON AND SURINAM 

Kirby (1869 : 362) makes the following comment about Zschach's work: 'Zschach 
quotes Linnaeus and Fabricius, but not Cramer or any other authors; in fact, he 
would seem to have been quite unacquainted with Cramer's work, as it will be 
noticed that he redescribes several species figured in Pap. Exot.' 

As dealt with here, four of the species named by Gmelin from Zschach's descrip- 
tions are strictly synonymous with Cramer or Stoll names, and five likewise with 
Linnaean names. At first sight it may appear odd that such a high proportion 
of synonyms should be created at a time when only a fraction of the exotic fauna 
had been described. The answer lies in the limited provenance of much of the 
early material. I estimate that at least 18 of the 25 species described by Zschach 
were either from Surinam or Ambon, both major sources for species described by 
Linnaeus, Cramer, Stoll and others during the latter part of the 18th century. 
This is turn may be explained by the activities of the Dutch, who from the 16th 
century had merchants and soldiers travelling the world. These people often 
brought back 'trophies', notably from their colonies Ceylon, Java, Surinam and 
Ambon, and thus large quantities of natural history specimens had already accumu- 
lated from these places by the middle of the 18th century. Ambon (or Amboina) 
in particular, despite its small size, was a principal source, no doubt due to its 
importance as a Dutch military headquarters. 

With regard to the synonymy of the Gmelin butterfly names, relevant information 
concerning the Indo-Australian species described by Linnaeus is presented in a 
series of papers by Corbet (1941a; 19416; 1942; 1945; 1949)- No similar work 
exists for the many species described by Cramer and Stoll; the need has not been 
so acute due to the quality of their original illustrations. So far as I can judge, it 
has generally been assumed that the Cramer and Stoll specimens, with the possible 
exception of some material in Leiden (see below), have been lost or destroyed. 
The following notes relate to the realization in recent years that a percentage of 
the Cramer material apparently survives among material now preserved in the 
British Museum (Natural History). 

The text of De Uitlandsche Kappelen by Cramer, and as continued by Stoll, bears a 
number of references to the 'rich cabinet of C. van Lennep' and also that of J. C. 
Sylvius van Lennep; (Cramer, 1775: [v]). According to Horn & Kahle (1935 '.47; 1936: 
287), the collection of Pieter Cramer passed via C. van Lennep (or one 'van Lennep') 
to either A. J. van Eyndhoven (whose collections were auctioned in Rotterdam 
in 1861) or C. V. Felder. The collection of the latter passed to Walter Rothschild, 
and thus finally to the BMNH. The Rothschild collection contains a considerable 
number of specimens, ex Felder, which appear to be Cramer, or, at least, van Lennep 



56 R. I. VANE-WRIGHT 

material. Many of these bear characteristic labels, with the specific name as given 
in Cramer written in seraph italic capitals, together with a reference to the appropri- 
ate plate. The style of the script bears a very close resemblance to the caligraphy 
employed in the frontispiece to the plates of Cramer (1775). Some of these specimens 
also bear circular blue labels of the form employed by Felder, and which normally 
refer to a country of origin, plus the designation 'Lenep' or 'Lenip'. It thus seems 
that the source of any likely Cramer material in the Felder collection was indeed 
one or both of the little-known van Lenneps, and that any specimen bearing reference 
to 'Lenep', etc., whether it carries a 'Cramer label' or not, should be examined very 
closely. The BMNH also possesses the original pattern plates for the works of Cramer 
and Stoll, which facilitates the necessary comparisons. Experience has shown that 
while a number of specimens with 'Cramer labels' may reasonably be accepted as 
part of the original type-series, others do not correspond closely enough to the original 
illustrations at an individual level to be accepted unhesitatingly as original type- 
material. Similarly, some specimens without the Cramer label, but with a Lennep 
label, seem almost certainly to be type-specimens. Each case has to be treated 
individually. Eventually it may be possible to produce a catalogue of all such 
butterfly specimens in the BMNH, cross-checked with material in Leiden (see below). 

An as yet unsolved mystery concerns the inclusion of a 'Cramer label' of the 
type under discussion in Horn & Kahle (1936 : pi. 26, fig. 4). A label of the supposed 
Cramer type is clearly shown in the photograph, but the caption ascribes it to 
'Halbton'. There is no other reference to Halbton in Horn & Kahle, and in fact 
I have been quite unable to discover any entomologist of such a name. I suggest 
that Mr Halbton, whose name translated into English means half-tone or semi-tone, 
is chimerical. It seems to me most likely that the 'Cramer labels' were written 
either by Cramer, Stoll or C. van Lennep. 

With respect to Cramer types in Leiden, Dr R. de Jong (personal communication) 
informs me that there appear to be none which certainly pertain to the Cramer 
and Stoll names discussed in the present paper. Dr de Jong also indicates that at 
present there is no general account of the Cramer material in Leiden. The probable 
Cramer specimens in the RNH evidently stem from the collections of Raye de 
Breukelerwaert, M. van der Meulen and van Lennep. Other Cramer material 
may exist in the Calkoen collection. According to Dr de Jong it is almost impossible 
to find any particular specimen which perfectly agrees with the figures given in 
Cramer and Stoll. In my opinion the illustrations in De Uitlandsche Kappelen 
are both somewhat stylised and idealised, but in a very subtle manner. 

SUMMARY OF TYPE INFORMATION 

Gmelin names. The following names described by Gmelin into the genus 
Papilio are dealt with in this paper: 

aethiops* , affinis*, annulatus* , argyrios, barbarus, bicolor*, bifasciatus, bipunctatus, 
chrysopterus* , claviger*, discors*, exoticus*, fenestratus,fuliginosus*,hexopthalmos*, 
hyalinus, lacteolus*, leo, leticostictos* , lunulatus, marinus*, oculatus, philetes, 
strigosus*, suipitia and vidua. 



BUTTERFLIES NAMED BY GMELIN 57 

Those names for which lectotypes have been designated are marked with an 
asterisk. In addition neotypes have been designated for bipunctatus, fenestrates 
and hyalinus. 

Type-material of other authors. Lectotype specimens have been designated 
in the present work for a number of nominal species described by authors other 
than Gmelin. These are listed alphabetically by author below, in their original 
combinations: 

Euplaea aglidice Boisduval, Danais leopardus Butler, Pyrrhopyga hospita Butler, 
Papilio calisto Cramer, Papilio itys Cramer, Eupioea pasithea Felder & Felder, 
Danaida juventa ogylla Fruhstorfer, Danaida limniace mutina Fruhstorfer, 
Delias dorimene avenda Fruhstorfer, Danais meganire Godart, Danais prothoe 
Godart, Pieris philyra Godart, Radena buruensis Holland, Tenaris nysa Hiibner, 
Papilio cenens Linnaeus, Papilio hecabe Linnaeus, Papilio pandarus Linnaeus, 
Papilio pipleis Linnaeus, Papilio urania Linnaeus, Cecropterus zonilis Mabille, 
Salpinx oculata Moore, Pyrrhopyga ulixes Plotz and Papilio cupentus Stoll. 

acknowledgements 

It is a pleasure to record my special thanks to Dr Colm O'Riordan, of the National 
Museum of Ireland, Dublin, for the generous help and assistance he has afforded 
me during the preparation of this work, both in London and in Dublin. I also 
wish to record my special debt to Dr H. K. Clench, Pittsburgh, Dr R. de Jong, 
Leiden, and Lt-Col. C. F. Cowan, who likewise aided with the substance of the 
paper. In particular, the sections on Gmelin's numbering system, and the problem 
of Papilio barbarus are almost entirely due to Lt-Col. Cowan. My thanks are due 
to Drs P. Viette, Paris, and H. J. Hannemann, Berlin, for making certain type- 
specimens available for study, Dr F. Martin Brown for photographs of Radena 
buriiensis types, and Mr E. C. Pelham-Clinton, Edinburgh, for photographs of 
certain Godart specimens. My colleague, Mr P. J. P. Whitehead kindly read draft 
sections of the manuscript, as did Mr R. B. Freeman, University College, London, 
Colm O'Riordan and Charles Cowan. I am further indebted to many of my col- 
leagues for help with particular points, including P. R. Ackery, B. J. Clifton, B. H. 
Cogan, R. W. Crosskey, D. S. Fletcher, P. Gilbert, P. M. Hammond, D. Hollis, 
T. G. Howarth, C. F. Huggins, I. W. B. Nye, A. C. Pont, D. R. Ragge, K. S. O. 
Sattler, R. L. Smiles, K. G. V. Smith, W.~H. T. Tarns, A. Watson and P. E. S. 
Whalley. My wife, Rosemary, kindly aided me with the preparation of notes 
and colour photographs of the Gmelin butterflies in Dublin. Messers P. A. Richens 
and P. V. York prepared the original photographs for the plates. Finally, I thank 
the Trustees of the British Museum (Natural History) for granting time and financial 
assistance for me to study the collections in Dublin. 

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1787. Mantissa Insectorum sistens Species nuper detectas 2, [ii] + 382 pp. Hafniae. 

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Herbst, J. F. W. 1792. Fortsetzung der Tagf alter. Vierte Familie. Parnassier. Natur sy- 
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Berlin. 



60 R. I. VANE-WRIGHT 

Herbst, J. F. W. 1973. Fortsetzung der Tagfalter. Sechste Familie. Rundfliigel mitbunten 

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Ent. Beih. Berl.-Dahlem 2 : 1-158, 16 pis; 3 : 161-296, 10 pis. 
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152. 

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BUTTERFLIES NAMED BY GMELIN 61 

Mabille, P. 1883. Description d'Hesperies. Annls Soc. ent. Belg. 27 : li-lxxviii. 

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Moschler, H. B. 1877. Beitrage zur Schmetterlings-Fauna von Surinam. Verb,, zool.-bot. 
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Plotz, C. 1879. Die Hesperiinen-Gattung Pyrrhopyga und ihre Arten. Stettin, ent. Ztg 
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Praeger, R. L. 1949. Some Irish naturalists, 208 pp., frontispiece -+- 18 pis. Dundalk. 
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Rober, J. 1 891. Beitrage zur Kenntniss der Indo-Australischen Lepidopterenfauna. 

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Amstelaedami. 
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Bayern. 



62 R. I. VANE-WRIGHT 

Staudinger, O. & Rebel, H. 1901. Catalog der Lepidopteren des Palaearctischen Faunenge- 

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Lipsiae. 



INDEX 



63 



BUTTERFLIES NAMED BY GMELIN 



Synonyms are in italics. 



acandra, 35 
acutipennis , 52 
aethiops, 49 
affinis, 35, 36 
ageleis, 34 
aglidice, 41, 42 
Alcides, 29 
Alcidis, 30 
a/ca, 36 

amboinensis , 42 
amyntas, 53, 54 
ancaeus, 25 
annulatus, 47 
anthyparete, 32, 33 
Antirrhea, 24, 46 
anzuletta, 44 
argyrios, 23, 29 
avenda, 34 

barbarus, 25, 53 
bi color, 31 
bifasciatus, 24, 46 
bipunctatus, 51 
bisaltide, 43 
brevipennis, 50 
buruensis, 38, 39 

cacica, 43 
caencus, 32 
calisto, 45, 46 
celeno, 54 
celmus, 32 
cenea, 33 
ceneus, 32, 33 
chrysopterus, 34, 35 
claviger, 38 
coridon, 54 
crameri, 42, 43 
crocale, 32 
cupentus, 47, 48 

deois, 45 
discors, 32 
diversa, 35 
dorimene, 33, 34 
dracontis, 30 
duplex, 40 



elissa, 47 
ellops, 43 
exoticus, 36, 37 

fenestrates, 52 
fuliginosus, 34 

hecabe. 34, 35 
hephaestos, 49 
hephaeslus, 49 
hewitsoni (Hypolimnas), 45 
hewitsonii (Jemadia), 50 
hexopthalmos, 42 
hospita, 49, 50 
hyalinus, 30 
hyperantus, 47 

isidora, 43 
idse, 31 
itylus, 43 
itys, 43, 44, 45 

jaira, 39, 40, 41 
jairus, 39, 40, 41 
juventa, 38 

lacteolus, 45 
lena, 31 
leo, 53, 54 
leopardus, 37 
leopoldina, 44 
leucostictos, 41 
leskii, 23, 29 
limniace, 36 
lisetta, 50 
lunulatus, 48, 54 

marinus, 39, 40 
meganire, 38, 39 
menechmus, 50 
murinus, 39 
mutina, 37, 38 

neis, 51 

nemertes, 41, 42 
nox, 40, 41 

nysa, 40 

oculata, 41, 42 
oculatus, 48 



6 4 



R. I. VANE-WRIGHT 



ogylla, 38, 39 
orontes (Alcides), 29 
orontes (Autochton), 51, 52 
orontiaria, 29 

pandarus, 45, 46 
pandemos, 40 
pasithea, 41, 42 
phaenareta, 35, 36 
philetes, 54 
philiasus, 53 
philyra, 32, 33 
pipleis, 45, 46 
pirithous, 25, 53 
plexaris, 32, 33 
polibete, 43 
prothoe, 36 
psenditys, 43 

russeus, 44 



sandya, 54 
sectator, 23, 26, 31 
strigosa, 43, 44 
strigosus, 43, 44 
sulpitia, 46 

talus, 54 
telicanus, 53 

ulixes, 49, 50 
urania, 39, 41 

vidua, 47 
vulpecula, 44 
vulpina, 44 

Zaretis, 44 
zethus, 43 
zonilis, 51, 52 



R. I. Vane-Wright, B.Sc. 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 



PLATE i 

Fig. i. Title page of Museum N. G. Leskeanum, Pars Entomologica, by I. I. Zschach (1788). 
Fig. 2. Title page of Museum Leskeanum, Regnum Animale, volume 1, byD. L. G. Karsten 

(1789). 

Fig. 3. Page from Karsten (1789) prefacing Class 5, the Insecta, which comprises an inclusion 
of the Pars Entomologica (Zschach, 1788). 

Fig. 4. Title page from O'Reilly's Dublin Society catalogue (1813). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 2 



PLATE 1 



MUSEUM 

N. G. LESKEANUM 

PARS ENTOMOLOGICA 

A D 

SYSTF.MA ENTOMOLOGIAE CI.. FABRICU 

CKOINATA 

I. F. ZSCHACH1I, M. Bacc. 



CL'M TAB, AEN. fICTIS. 

__. !■■ ■ 1. . . if J rrxa^x= 

I. I P S I A B, 

IN BIELIOPOLIO I. G. MULLER1ANO. 

1788. 



1 J /£ ~*\ 



[N] 



Catalogue 

OF Tlir SUBJECTS OF 

ATURAL HISTORY, 

MUSEUM 

THE RIGHT HON. AND HON. 

THE DUBLIN SOCIETY, 

ftTSTEBUTICALLf ARRANGED) 

THE ANTIQUITIES, #c. 



MVSEVM 

LESKEANVM 



REGNVM ANIMALE 



6.VOD 
ORDINK SYSTEM ATICO 

DtSPOSViT ATQVE DEfCRlPSIT 

D. L. GVSTAVV'S KARSTEN, 



C I E T. NAT. 



" A 1 1 RB, tOUALII. 



V L I. 

C « in IX. i colli bus p 1 £1 i s. 



L I P S I A E, 

SVMPTIBVS HAEREDVM I. C. MVLLERr. 
■ - 8 9 



CLASSIS V. 

I N S E C T A. 

C U R A J. J. Z S C H A C H 1 I. 



Hicc cljflis a Claiifl" J. J. 2fer.axhio Mtd. Rj.r. elaborata, urn 
ante annum tt quod excnrrit Inb tituJu: Muren.;! N. C. 
I.wkcanum. Pari cnlomotoeica. adfylfcma cntoinoloai,,!! 
Ct. Fabricif ordinals g inaf, in uu)MapolioMulIcriajtoty»i< 
cxprc-naell. Qiiic cMoruoiogu-a rolirctiD, cum rtngula- 
rem oollrfluri. curam cxpoftat, f..n>ir,s a reliquo inulio 
fnuntia. natural- lirauuri, mius imprimis inrtrllt, 
ittanj pofljdcrx , fcparariui nivtndetur. 



PLATE 2 



Fig. 5. Page 87 of Zschach (1788), showing the first seven species of Papilio. The 
unnamed no. 6 was subsequently named argyrios by Gmelin (1790) and finally leskii by O'Reilly 
(1813); it is now considered to represent the moth species Alcides orontes (L). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 2 PLATE 2 



PARS ENTOMOLOGICA. g 7 

C L AS S I S VI. 

G h O S S A T A. 



A 



130. PAPILIO. 

* j. * Equites Troes. 

I. P. E. T. Deiphobus. F. S. E. T. 8- in Afict. 
3. P>E.T. Hector. F. S. £. T. 4. in Indus. 

3. P. E.T. Pammon. F. S. £. T.13. inAfia. 

4. P. E.T. Polydamas. F.S. £. 7". ££. in America. 

5. P. E. T. Aeneas. F. S. E. T. 23. in India. 

* 2. * Equites Achivl 

6. P.E-. A. '' alis dentatis, fupra nigris, anticis fafciis duabus' 
argentcis, poftica nebulofa, fabtus coeruleo-argcnteis,fa- 

.. .fciis duabus > apice niargineque poftico unitis, fufcis, 
ftriae transverfae, coeruleo-argenteac verfus bafin, utrin- 
que ad marginem craffiorem, alis pofticis dentatis, dente 
intermedio, rejiq'uis longiore, fupra fafcia latiori argen- 
tea, maciijisque 7 coeruleo-argenteis, lineola transverfa 
interrupt^ ,ad marginem pofticum ; fubtus coeruleo-ar-? 
A genteae, in medio maculis tribus transverfis aeneis, una- 
quc remota ad marginem internum, fafciisque duabus ni- 
gris, prima undata i altera dentata-, margineque poftico 
nigro ? albo macnlato, inter fafciam pofticam ct margi- 
nem maculae coeruleo-argenteae, anticis lineolis nigris 
. interruptae, maculae trcs dentatae", nigrac, ad marginem 
internum verfus anguium anij abdomen fubtus luteum, 
Exoticus. 

7. P. E. A. Lcilus. F. S. E. A. jr K in Americct. 



PLATE 3 

Fig. 6. Entry on page 89 of Zschach (1788) for the unnamed 'Papilio 46c'. 

Fig. 7. The description of Papilio claviger Gmelin (1790 : 2289) as it appears in the 13th 
Sy sterna Naturae (cf. fig. 6). 

Fig. 8. Part of p. 75 of O'Reilly (1813); compare the entry for Papilio 46c with figs 6, 7. 

Fig. 9. The entry by Kirby (1869 : 358), in his work on the Gmelin names, for Papilio 
claviger. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 2 



PLATE 3 



46. c. P.D. f. ® alis ftibintegris, fufcls, maculis purjctisque al- 
bis, hyalinij, bafi artticis longis, polticis radiatis, clava- 
tis, margimbus pofticit duplici feric punctor::inalborutn, 
fubfus conculores. Exoticus. 



claviger. 887 P- alis ftibintegris fufcis : maculis pun&isque albis hyali- 
111s bjlt atiterionbus lungis, polterloiibuj radiatis clava- 
tis, marginibus polteriocibus duplici ierie punctorum al- 
. boi 11m. • Muf. Let/t. />. 89. ». 46. c. 
Habitat extra Europain. 



75 



8 



46° P 

47 

48 

49 

50 

51 

52 

53 

54 

55 

56 

57 



D. F. Claviger Club-spotted Butterfly 

Murinus Mouse-couloured ditto 

Leucostictos White-spotted ditto 
HexophthalmosSix-dull-eyed ditto 



(.'hi. 1 mxus 

Xanthus 

Antiochus 

Plexippus 

ChrysippUS 

Strigosus 

Hyperunthus 

Eribotes 



Choriuaeus ditto 
Xanthus ditto 
Antiochus ditto 
Plexippus ditto 
Chrysippus ditto 
Streaked ditto 
Brown Argus ditto 
Eribotes ditto 



7. Nympfides Gemmati. 



India 
lb. 
lb. 
lb. 

Surinam 

Asia 

China 

America 

India 

lb. 

Europe 

India 



58 P. N. G. Iris 



58 b 
59 
60 



Lacteoius 
Bif'asciatus 
CEuone 



Emperor of the Woods or 

Purple Highflyer Europe 
Milk-dropped ditto 
Banded ditto 



CEnone ditto 



India 

lb. 

Asia 



9. Papilio claviger, Gmel. p. 2289. 

P. D. F. alis subintegris, fuscis, maculis punctisque 
albis, hyalinis, basi anticis longis, posticis radiatis, cla- 
vatis, marginibus posticis duplici serie punctorum alborum, 
subtus concolores. Exoticus. (Zschach, p. 89, n. 46c.) 

The apparent type of this insect is very similar to, but 
not quite identical with, our specimen of Danaus Juventa, 
Cram. 



PLATE 4 

Fig. io. The 'Advertisement' by Bernard O'Reilly which prefaces his 1813 catalogue of 
the Dublin Society collection. 

Fig. 11. Part of p. 73 of O'Reilly (1813), showing the commencement of the butterflies 
and including the introduction of the name Papilio leskii (cf. PI. 2, fig. 5). 

Fig. 12. Entry in the British Museum (Natural History) library catalogue (1910) for the 
book on Greenland, etc., by Bernard O'Reilly (1818). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 2 



PLATE 4 



ADVERTISEJLEtfT. 



10 



TV yy?0OM( Catd.^ti rilubUi ut aaa iw, la n/ttfaulk tlau^t. 
tum. a jiral auatacr of ii.bjaU a/ S.inral fultrj, la '*< ZnaUg.J 

TV &•*» ('. fi*TaL'f. lid of l.l.HJCV, tor turpi*, ktrog ra lot 
Chu InaccM, uii.-A o DM^«Mh fa F*Miriu». m/* o/artW 
tfmattaa, aaJ.r »*' miiJwujf j?" Cunaa, ay «Ah-^ (Aa G«mi Cancer, i« 
*ilkir**m fro* ikt Araaa'/Lov^ci, awtfiA* A(Oju./r.i«iun, 
yi> lie <3aM Varan*, aaaar a wi War CfvuLtocn. 

£ai-A arl«/. u a; ray"* in lit fJtt*=n. or Jtr. .« — firrf, l*r ai£urri 
wai-A outraf^tt s«A ihr m..nlrr.ofij,.lt.>'kt S-il.jo.1 i* fall Miucm $*• 
raaaVy, l»' Ccorri. aad tlmjil aaatr ra ia/". IhrJlt. iKt r.flrW) 
n-.V.J naw '/Mr m ,o Ev.LJ. /W*«y, lot Jbbun V /'''<-< 

a.irr I'nucJ. * 

The MiKiil^aeatt sarf a/iAe f„i.n>...a imUtt a ta-i.ty g^nnaM 
Aotij.r nVautaj < aZlir /i./io» OmaaiaaU and Arvt, .elk many tngrti ' i 
wii ■/ Jrt. 

Ha lodfj a* (A; rarf eVrad. (a lAa flj^raai /Wa ta Iht .Waiaam, fa 
Wara Ua C. -c. . .1 refrrt. 

BEBXAHD aSHlLLi. 



Order VI. Glossala. 

Mouth with palpee, and a spiral trunk. 
Papilio. 



1 Equites Troes — Trojans. 



1 P. E. T. Deiphobus 
'2 Hector 

3 Pammon 

* Polydamas 

.5 yEneas 



Deiphobus Butterfly 
Hector ditto 
Pami.ton ditto 
Polydamas ditto 
Eneas ditto 



6P.E.A 
7 
8 
9 
10 



2. Equites Ackivi — Greeks. 



Asia 

India 

Asia 

America 

India 



Leskii Leskean ditto India 

Leilus Leilus ditto America 

Podalirius Scarce Swallow-tail ditto Europe 

Machaon Swallow-tail ditto lb. 

Dcmoteus Demoleus ditto India 



11 



12 



O'REILLY (Bernard) Greenland, the adjacent seas, 
and the North- West Passage to the Pacific Ocean, 
illustrated in a voyage to Davis's Strait during the 
summer of 1817, &c. pp. vi, 293 : IS pis., 3 maps. 

4°. London, 1818. 

According to the " London Quarterly Review" (Vol. XIX, pp. 208-214) 
this is " one of the most bare-faced attempts at imposition . . . The very 
small portion of his Greenland which is not absolute nonsense is either 
fiction or dov nright falsehood." 



PLATE 5 

Figs 13, 14. Entries for Danaus meganire (p. 5) and D. claviger (p. 8) in Kirby's catalogue 
(1871). 

Fig. 15. Entry on p. 691 of the Supplement (1877) to Kirby's catalogue, showing the synony- 
my of claviger and meganire. 

Fig. 16. The cryptic entry '19 = no. 29', which is the original synonymy of claviger with 
meganire, appearing on p. 639 of the Appendix to Kirby's catalogue (1871). 

Figs 17, 18. Examples of manuscript entries by W. F. Kirby in the interleaved copy of 
Zschach (1788) in the National Museum of Ireland. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 2 PLATE 5 



19. D.Meganira, Godt. Enc. M6th.IX. p. 192. n. 51. Ins. Ind. 
TO (1819); Blanch. Voy. Pol Sud, p. 387. t. 2. 

' ° f. 4. (1853.) 



29.? D. Claviger, Gmel. (Pap. C.J Syst Nat I. 5. p. 

M2289. 11. 887.(1788— 1793); Pap. — . Zsckach, 
Mus. Lesk. Ent p. 89. n. 46 e. (1788.) 



29. D. Claviger, Gmel. (p. 8.) . 

]C D. Meganira, Godt. & D. Sobrina, Boisd. (nos. 19 5 21, 

l ° V- 5.) 



76 19 = no- 29 ; 30 = no. 15. 



'jr. $-/l. z^? a -» ^/ J • 



• o £> 7.77 ?/ ; 6 /••; .0 



PLATE 6 

Fig. 19. Pierella hyalinus hyalinus (Gmelin), neotype $, Surinam, Lenep; this specimen 
is the one believed to have been figured by Cramer [1780 : 5, pi. 291, figs A, B] as Papilio Una 
Linnaeus (fore wing length 37-5 mm; BMNH). 

Fig. 20. Danus juventa claviger (Gmelin), <$, Indes Orientales [? Ambon]; lectotype of 
Danais meganire Godart (fore wing length 41 mm; MNHN). 

Figs 21, 22. Danaus limniace exoticus (Gmelin). (21) lectotype of Danaida limniace 
mutina Fruhstorfer, $, Ceylon 1889, Fruhstorfer (fore wing length 45mm; BMNH); (22) 
lectotype of Danaus leopardus Butler, £, N. India (fore wing length 49 mm; BMNH). 

Fig. 23. Jemadia hospita hospita (Butler),^, 'Surinam' [false locality?]; lectotype oiPyrrho- 
pyga ulixes Plotz (fore wing length 27-5 mm; MNHU). 

Fig. 24. Jemadia hospita hephaestos (Plotz), holotype $, Surinam (fore wing length 30 mm; 
MNHU). 

Fig. 25. Jemadia lisetta Mabille & Boullet, g syntype, Peru (fore wing length 27 mm; 
MNHN). 

Fig. 26. Autochton bipunctatus (Gmelin), neotype <$, French Guiana (fore wing length 21 mm; 
BMNH). 

Fig. 27. Ablepsis fenestratus (Gmelin), neotype $, French Guiana, Nouveau Chantier, 
Le Moult; this specimen is also the holotype of Telemiades acutipennis Mabille & Boullet (fore 
wing length 21 mm; MNHN). 



Bull. Br. Mus. not. Hist. (Ent.) 32, 2 
V / 




PLATE 6 












27 



ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 

1965- £3-25- 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3-i5. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Chora (Lepidoptera: Geometridae) . Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae) . Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£19- 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. 
February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text-figures. December, 1973. £9-55- 

printed by Un win Brothers Limited the gresham press old woking surrey England 



JUNI975 

RE VISIONAL NOTES ON AFRICAN ^ 
CHARAXES, PALLA AND EUXANTHE 
(LEPIDOPTERA : NYMPHALIDAE) 

PART X 



V. G. L. van SOMEREN 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 3 

LONDON : 1975 



REVISIONAL NOTES ON 

AFRICAN CHARAXES, PALLA AND EUXANTHE 

(LEPIDOPTERA : NYMPHALIDAE) 

PART X 




BY 

VICTOR GURNER LOGAN van SOMEREN 

The Sanctuary, Ngong, Karen, Kenya 



Pp. 65-136; 19 Plates; 9 Text-figures; 3 Maps 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 3 

LONDON : 1975 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 3 of the Entomological 
series. The abbreviated titles of periodicals cited follow 
those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent.) 

ISSN 0007-6431 



© Trustees of the British Museum (Natural History), 1975 






TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 5 June, 1975 Price £13.00 



REVISIONAL NOTES ON 

AFRICAN CHARAXES, PALLA AND EUXANTHE 

(LEPIDOPTERA : NYMPHALIDAE) 



PART X 



By V. G. L. van SOMEREN 



CONTENTS 

Page 

Synopsis ........... 67 

The Palla complex ......... 68 

Palla decius (Cramer) ........ 69 

Palla ussheri (Butler) and its subspecies .... 70 

Palla violinitens (Crowley) and its subspecies .... 73 

Palla publius Staudinger and its subspecies .... 75 

Systematic List ......... 77 

The Euxanthe complex ......... 78 

Euxanthe eurinome (Cramer) and its subspecies .... 79 

Euxanthe crossleyi (Ward) and its subspecies .... 83 

Euxanthe wakefieldi (Ward) ....... 85 

Euxanthe madagascariensis (Lucas) ...... 86 

Euxanthe (Hypomelaena) trajanus (Ward) and its subspecies . 87 

Euxanthe {Hypomelaena) tiberius Grose-Smith and its subspecies 90 

Systematic List ......... 92 

A brief historical reference ....... 94 

Corrigenda to parts I-IX ........ 95 

Addenda ........... 102 

further descriptions and notes on species and subspecies of 

Charaxes .......... 103 

a further note on the 'Black' Charaxes complex . . . 107 

SOME REPLACEMENT NAMES AND ADDITIONAL INFORMATION ON 

CERTAIN TAXA ......... Io8 

Synoptic list of African Charaxes, Euxanthe and Palla . . . no 

Acknowledgements . . . . . . . . . 118 

References for parts I-X . . . ..... 119 

Appendix ........... 127 

Index to parts I-X ......... 127 

Maps ............ 133 



SYNOPSIS 

This final part of the series deals with the Palla and Euxanthe complexes, each of which, 
following Rydon (1971 : 231), is regarded as constituting a separate subfamily. Three new 
subspecies are described in Palla and one in Euxanthe. A synoptic list of all the species and 
subspecies dealt with in this series is given, together with a complete list of references and a 
complete index to all ten parts; there is also a list of corrigenda to the earlier parts. In the 
Addenda section dealing with Charaxes, three new subspecies, one form and one aberration 



68 V. G. L. van SOMEREN 

are described, nine taxa of the 'black' Charaxes complex are raised to specific status and five 
new names are proposed to replace junior homonyms. 

THE PALLA COMPLEX 

In this complex there is a group of characteristic species, the males of which are 
confusingly alike, all conforming to a similar pattern on the upperside with black 
fore and hind wings crossed by conspicuous bars, white in the former and white 
and tawny orange-brown in the latter. The undersides also exhibit a very similar 
ground colour on both fore and hind wings, crossed by a white bar which passes 
through both. On the other hand, the females are more distinctive, one species 
having a male-like female, the others having females which are more sexually 
dimorphic. 

Members of this complex are usually placed in the subfamily Charaxinae, but 
in view of the distinctive characters of their ova, larvae and pupae, which differ 
markedly from those of Charaxes, the restricted food-plants, which belong to the 
Convolvulaceae, the different habits and, moreover, the genitalia, which are 
distinctive and quite unlike those of Charaxes, I agree that they should be placed 
in a separate subfamily, the Pallinae, the name proposed by Dr A. H. B. Rydon 
(1971 : 230). 

Aurivillius (1912) has pointed out that there are characters in the venation, 
especially in the hind wing, and mentions that the middle and hind tibiae are 
without spines above. These may be considered merely generic characters. 

Schultze (1917 : 593~595) also has some cogent remarks to make on the genus. 

For an account of the early stages see Schultze (1916 : 126) for ova and larvae, 
van Someren & van Someren (1926 : 350, pi. 77, figs 1, 2), van Someren & Rogers 
(1930 : 31, pi. 106) and Rydon (1971), also the line drawings which accompany 
these notes (see Text-figs 7-9, p. 79). 

When considering the problem of zoogeographic variation, one must constantly 
keep in mind the fact that one is dealing with species distributed throughout several 
more or less well defined ecological areas which are as follows. 

Occidental Africa, comprising Senegal, Guinea, Sierra Leone, Ivory Coast, 
Liberia, Ghana, to western Nigeria. 

Eastern Nigeria, east of the Cross River, Cameroun, Central African Republic, 
Zaire, Congo (Brazzaville), N. Angola (a subdivision of this includes the south- 
western corner of Cameroun, Guinea and Gabon). 

The Congo Basin, including the Kasai district to eastern Zaire, western Kivu 
district, Stanleyville, Nawamba-Beni-Irumu, west of the Semliki River. 

The Katanga district. 

The Central Rift areas: Rwanda, Burundi, S.W. Uganda, especially the Kigezi 
country, the Bwamba VaUey west of Ruwenzori, and the districts of western 
Uganda. The eastern subdivision includes central Uganda and N. W. Kenya, 
Elgon area. 

The north-western corner of Tanzania including the Kigoma district to the 
north-east of Lake Tanganyika. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



69 



With the above in mind, one may attempt to solve the Palla problem. The species 
involved are: Palla decius (Cramer, 1777), P. ussheri (Butler, 1870) P. violinitens 
(Crowley, 1890) and P. publius Staudinger, 1892. 



Palla decius (Cramer) 
(PI. 1, figs 1-4; Map 2) 

Papilio decius Cramer, 1777 : 26, pi. 114, figs a, b. 

Palla decius (Cramer) Kirby, 1871 : 273. 

Palla decius f. Sagittarius Rousseau-Decelle, 1934 : 235. 

Male. Fore wing length 38 mm. Shape falcate, but apex rounded; the outer margin 
concave at 4-5; hind angle projecting but rounded. Upperside. Fore wing, ground colour 
black, wing crossed by a white bar commencing just beyond the cell, 5 mm at costa and 
expanding gradually through spaces 2-3 and reaching the hind margin at mid point where it is 
7 mm wide; its inner border with some blue scaling most conspicuous at end of cell and bases 
of 2-1&. Hind wing, ground colour black on border, slightly browner at base and inner fold 
where there is a whitish streak above the anal angle, the disc of the wing crossed by a white 
bar 5 mm wide at costa, reaching the upper part of the cell, then shading to orange and orange- 
tawny and expanding distally to the hind angle to reach vein 4. There is a conspicuous black 
rounded spot on submargin at vein 2 and an orange spot with white-centred black in the sub- 
margin above the orange-tawny tail at end of vein 4, with traces of a spot in cellule above. 
Edge of wing slightly undulating; anal angle squared, but rounded at tip. Underside. Fore 
wing, basal area rich brownish black, distally flanked by a conspicuous white bar, more or 
less as above; the distal portion of the wing with obscure silvery white ground in postdiscal 
zone, crossed by fine black lines in decreasing distance apart until on the border where the 
ground colour is uniform brownish black. Hind wing, basal area and greater part of inner 





Figs i, 2. Euxanthe, $ genitalia. 1, E. wakefieldi (Ward) (Tanzania: Amani), slide no. 
R. 337; 2, E. tiberius Grose-Smith (Kenya: Shimba Hills), slide no. R. 338. (A. H. B. Rydon 
del.) 



70 V. G. L. van SOMEREN 

fold deep brownish black, finely lined on the inner fold, and sharply defined from the white 
bar which on its distal side and the border is silvery white with fine cross lines of increasing 
density toward the border, with some silvery rays at the edge above the tail and at anal angle. 
Margin rusty brown with whitish lunules proximally, carrying blue-black dots, enlarged to an 
ocellus in space 2, and some silvery scales at anal angle, but edge above the tail blackish 
brown. 

Female. Larger than the male, fore wing length 40 mm. Shape like that of the male, 
but slightly less falcate. Upperside. Fore wing, ground colour, base more brownish, distal 
portion also brownish. The white bar conspicuous as in the male but more curved on its 
inner edge, the upper portion of the bar shading to orange distally and on the outer edge, 
especially in spaces 4-3. Postdiscal zone with a series of tawny-orange spots, palest at costa 
and rather angled; spot in space 4 smaller and set in. Hind wing, base and upper part of 
inner fold brownish. Disc of wing crossed by a broad white bar of equal width, 4-5 mm wide, 
but tapering to above the hind angle. Border of wing brownish, with a series of pale rufous 
spots distally, from upper angle to tail; a marked ocellus with blue-black centre at 2; anal 
angle rufous with two white dots. Tail at end of vein 4 long and rounded at tip, 4-5 mm. 
Underside. Fore wing, basal area brown, edged black; a few dark lines margined in white in 
the cell, bordered distally by a whitish bar, sharply defined proximally but shading to buffy 
distally, the edge dyslegnic and merging into the greyish brown border which is lined with 
fine dark fines of increasing density, becoming brownish on the border. The disco-submarginal 
zone with a series of buffy ochreous spots corresponding to those above present, but rather 
obscured. Hind wing, basal area brown, similar spots to those of fore wing in disco-submarginal 
area but more lunate; a conspicuous ocellus at vein 2 with blue-centred black spot. Anal angle 
rufescent with two white dots; border of wing slightly rayed at upper angle and in the region 
of the tail; margin of wing rufescent with small whitish blue dots on admargin; edge brownish. 
The inner fold has fine dark lines. 

The underside of this species is more variegated than the majority of those in the group. 

Range. French Guinea, Sierra Leone, Congo (Brazzaville), Kasai and N. Angola. 



Palla ussheri (Butler) 

Valla ussheri ussheri (Butler) 

(PI. 1, figs 5-8; PI. 2, figs 9, 10, 14; Text-fig. 3; Map 2) 

Philognoma ussheri Butler, 1870 : 124. 
Palla ussheri (Butler) Kirby, 1871 : 273. 
Palla ussheri f. fevruginea Schultze, 1914 : 83. 

Male. Fore wing length 35 mm. Shape falcate but apex rounded; hind angle projecting 
slightly. Upperside. Fore wing, ground colour black, crossed by a conspicuous white bar, 
as in decius, but narrower throughout, the inner edge only slightly shaded with greyish, the 
portion in space \a is oblique on its inner side. Hind wing, ground colour brownish black at 
base shading to brownish on inner fold; border of wing black, widest at 4-5, then tapering to 
above the hind angle ; the intervening space at costa at vein 6 is white, then shading to orange 
and tawny-orange, clearly defined basad and extending to outer side of tail at vein 4, and on 
the inner border shading to brownish on the inner fold which has a pale spot at the anal angle; 
the space above the tail with black-centred orange spot and a more conspicuous black spot 
with white centre in space 2 of the submargin. Tail 4 mm long, slightly tapered, tip rounded 
and pale. Underside. Very similar to decius but rich brown of basal areas of both wings more 
curved on outer edge. Fore wing, cell with four black and white cross lines; the inner crossed 
by fine wavy black lines. The conspicuous white bar narrower, but expanded to 6 mm at the 



REVISIONAL NOTES ON AFRICAN CHARAXES 71 

hind margin, its outer edge flanked by a zone of fine black lines darkening to a series of obscure 
dark spots in the submarginal zone which extends from the subapex to the hind margin; border 
brownish. Hind wing with narrow white bar, gently curved, accentuated proximally by black, 
and on its distal side flanked by fine black lines gradually becoming closer and more dense 
toward the submargin, but with a pale streak along vein 4. The submargin brownish black 
with brownish marks in admargin with increasing black, white-centred dots, that in space 2 
forming a conspicuous ocellus with marked central spots and with a white line proximally; 
the anal angle more rufous with two white dots; edge brownish black. 

Female. Resembling female decius somewhat, but the colours brighter and stronger. 
Uppeyside. Fore wing, base dark brownish, paler along the inner fold; distal portion of wing 
blacker, the intervening white bar, strongly defined on its inner border, is shaded with orange 
and very faint wavy black lines distally, becoming less strong toward the hind border, 5 mm 
wide at costa and gradually widening to 1 1 mm at the hind margin. A conspicuous series of 
tawny orange spots present in the postdiscal submarginal line, small and pale at the costa, 
large in spaces 5-3 to the hind angle, the spot in 4 inset a little. Apex and border of wing 
brownish black. Hind wing, white bar slightly shaded with greyish proximally, and with 
orange distally, is 8 mm wide at the costa, slightly less at 4, then tapering to above the hind 
angle where there is a white streak. Distal portion of wing blackish, shading into the orange 
on the distal side of the white bar. In the submarginal area is a series of large orange spots, 
slightly paler proximally, the lower ones with purplish-centred dots developing into a 
conspicuous ocellus in 3; the anal angle with two white dots. Tail at end of vein 4 robust, 
rounded and pale at end, 5 mm long. Underside. Generally paler than in the male, the basal 
areas of both wings paler brown; fore wing, white bar outlined in black proximally, much 
narrower than above, being encroached upon by the greater extent of the wavy black lines, 
especially at the upper end. The submargin carries a series of confluent tawny-orange spots 
outlined in black at apical end, the two subapical spots whitish, partly obscured by the wavy 
lines; border of wing more uniform pale brownish. Hind wing with a restricted white bar 
sharply defined proximally, but reduced distally by the very dark lines which extend to the 
submargin where there is a series of tawny-orange spots, white-edged proximally and each 
bearing a white-centred black dot, strongly marked above the tail and represented in space 3 
by a conspicuous ocellus which is shaded with olive distally. The anal angle is orange-tawny, 
with two white dots. Pale rays are present in the border along veins 4 and 6. 

Range. Sierra Leone to west Nigeria, west of the Cross River. 

£ form dobelli Hall stat. n. 
(PI. 3, figs 19, 20; Map 2) 
Palla dobelli Hall, 1919 : 199. 

Described from a single female taken at Bitje, Cameroun, this specimen was 
placed as a form of Palla moderata Gaede, 1915, also from Cameroun, by Bryk 
(1939 : 547). It has also been placed under Palla ussheri and considered to have 
precedence over ussheri interposita Joicey & Talbot, 1925, described from Uganda. 

In my opinion dobelli is one of the female forms found amongst the intermediate 
aggregate, ranging from Cameroun to E. Congo (PL 2, figs 12, 13, 15). In this 
aggregate, the males are very similar to nominate ussheri, both above and below, 
but the pattern is less contrasting. The same applies in a general way to the females, 
where the variation is from a specimen very similar to the nominate form, to one 
in which the upperside pattern is similar to that of the type dobelli, the underside 
pattern also less strong than in the nominate form. 



7 2 



V. G. L. van SOMEREN 



Palla ussheri interposita Joicey & Talbot 
(PL 3, figs 18, 21-24; PL 4. fig- 2 5; Text-fig. 4; Map 2) 
Palla ussheri interposita Joicey & Talbot, 1925 : 646. 

Male. Fore wing length 38-41 mm; shape as in the nominate race. Upperside. Compared 
with nominate ussheri general coloration and pattern very similar, but the fore wing white 
bar, though narrow at the costa expands to 8 mm at the hind margin and is shaded more with 
greyish proximally. The basal dark area not so blackish. The hind wing band, mainly 
orange to tawny-orange, though paler toward the costa, is wider, the orange area is less defined 
on its proximal border and wider at distal end but more restricted at base of tail, leaving the 
orange spot above the tail free. The ocellus towards tornus larger; the two white spots at 
anal angle more distinct. Underside. Forewing very similar to nominate race but basal areas 
paler brown ; the hinder part of the white bar less expanded and the distal side of the bar more 
encroached upon by the wavy dark lines which, however, have satiny, obscure spots in the 
postdiscal line and in the subapex. Hind wing exhibits similar differences as on the fore wing, 
the satiny whitish rays at veins 4 and 6 more distinct, so also the submarginal silvery lunules ; 
the ocellus at the outer margin larger and more conspicuous, the ground colour being more 
olive-brown; the tail dark but orange-tipped. 

Female. Larger than the male, fore wing length 42 mm. Upperside. Coloration rather 
different to that of the nominate race, much less strongly patterned. The basal areas of both 
wing bluish grey, slightly darker in the cell. Fore wing, white band strongly shaded with orange, 
leaving only a triangular white patch on proximal side; the band is 12 mm wide at the outer 
margin, the distal orange shading into the area of the postdiscal zone, reducing the width of 










Figs 3-6. Palla, g genitalia. 3, P. ussheri ussheri (Butler) (Cameroun), slide no. R. 334; 
4, P. ussheri interposita Joicey & Talbot (Uganda), slide no. R. 333; 5, P. publius 
Staudinger (Bipindi), slide no. 335; 6, P. violinitens (Crowley) (West Africa), slide no. 
R. 336. (A. H. B. Rydon del.) 



REVISIONAL NOTES ON AFRICAN CHARAXES 73 

the border and also causing the submarginal orange spots to be obscured. Hind wing, the inner 
edge of the white band is straighter, but suffused with orange distally, thus reducing the width 
of the dark border on which the orange spots are less distinct except at the hind border where 
the ocellus is very conspicuous. Tail at end of vein 4, n mm long, rufous but pale tipped. 
Underside. The general appearance of the underside is very similar to that of the nominate 
female, the main differences are in the reduction of the dark bases; in interposita the black lines 
with white edges are more obvious on a paler ground. The white bars on fore and hind wings 
more restricted distally, due to the extension of the dark wavy lines proximad. The submarginal 
dark mark of fore wing more distinct and on the hind wing the ocellus is larger and very 
conspicuous. 

Range. Uganda, mainly central and east; less plentiful in western forests. A 
very worn specimen which is rather intermediate between the nominate subspecies 
and interposita was captured at Lunzuwa Falls, Mbala, Zambia on 8.^.1969 by 
Mr F. Scofield (teste M. N. Mitchell, pers. com. 1973). 



Palla violinitens (Crowley) 

Valla violinitens violinitens (Crowley) 

(PL 4, figs 26, 27; Text-fig. 6; Map 2) 

Philognoma violinitens Crowley, 1890 : 554. 
Palla violinitens (Crowley) Staudinger, 1892 : 266. 

Male. Fore wing length 35 mm; shape as in other species of Palla. Upperside. Fore 
wing base and distal portion black, crossed by a conspicuous white bar, narrow at costa, 3 mm 
and widening slightly to the hind margin, strongly edged with blue on proximal side, slightly 
on outer in la-lb. No other markings. Hind wing brownish at base, this colour extending 
into the inner fold, at the anal end of which is a whitish mark. Border blacker, separated 
from the base by a bluish white band sharply defined on inner edge, but merging into rufous 
orange below cell, which colour extends to the hind margin and along the tail, with a separate 
orange spot in the black border above the tail. Tail at end of vein 4 orange and pale tipped, 
6mm long. Underside. Fore wing base deep chocolate-brown; cell crossed by five black, 
white-edged, lines. White bar strong and narrow as above, defined by black on inner edge, 
flanked distally by fine dark lines on a greyish ground, so that the distal part of the wing appears 
darker than in other species. Hind wing base deep chocolate-brown, white bar narrow, 3 mm 
at costa, then of even width to above the anal angle, outwardly flanked by the dark wavy lines 
on a greyish ground. Border of wing with rufescent spots, distinct above and below the tail; 
ocellus with black spot on an olive ground, conspicuous, shaded with black proximally. 

Female. Larger than the male, fore wing length 45 mm, apex more rounded. Upperside. 
Fore wing basal area blackish brown, white band well defined, the distal edge with fine dark 
lines and so merging into the dark brownish black apex and outer border. The white band 
at the costa 7 mm wide, gradually expanding to 15 mm on the hind border; the dark border 
with somewhat angular white spots, that in space 4 small and set in. Hind wing basal area 
blackish brown extending to inner fold well above anal angle, white band very broad, 15 mm 
at costa to vein 4, then tapering to inner fold above the hind angle. Border of wing brownish 
black, merging into the white band by fine dark wavy lines ; a row of conspicuous white lunules 
in the submarginal zone bordered distally with rufous above and below the tail; the ocellus 
on hind margin conspicuously white-edged proximally, with large central black spot; mark 
on anal angle paler, carrying two white dots. Tail at vein 4 rufous with a dark edge, tip pale, 
10 mm long, but robust with rounded tip. Underside. Basal areas of both wings chocolate- 
brown, with black lines in the cells ; outer edge with blackish line defining the broad white band 



74 V. G. L. van SOMEREN 

proximally, and distally set off by fine black wavy lines of increasing density toward the darker 
border and in the hind wing with a pale ray along 4; the submargin with white lunules edged 
rufous distally, forming ocelli above and below the tail, that on the hind margin conspicuous 
with black spot on an olive ground, accentuated proximally by a black line, that on anal angle 
olive, with two white dots. 

Range. Ghana to the Central African Republic and Angola, where it intergrades 
with the next aggregate. 



Cline coniger (Butler) 

(PI. 4, figs 28-32; PI. 5, fig. 33) 

Char axes coniger Butler, 1896 : 403. 

Char axes coniger Butler; Gabriel, 1927 : 34. 

Usually referred to as merely a form, the name coniger Butler is here considered 
to be applicable to the large aggregate of the species found east of Cameroun. It 
will be noted that the type male came from Old Calabar and the associated female 
from 'Congo'. 

Male. Very similar to nominate race in size and coloration. Upperside. Forewing differing 
only in the greater width of the white bar with less strong blue on its proximal side. Hind 
wing, band also wider in its white upper portion, but the orange portion is more restricted. 
There is little difference on the underside except a corresponding difference in the width of 
the white band as noted on the upperside. 

Female. In this sex there is a corresponding difference in the width of the bars on the 
upper and under surfaces to that noted in the males. Whereas in the nominate race the 
submarginal row of pale spots on upperside of both wings is white, in coniger these spots are 
white or orange. 

Range. Cameroun, Central African Republic, Congo (Brazzaville) and Zaire. 



Palla violinitens bwamba subsp. n. 

(PI. 5, figs 35-37; Map 2) 

Male. Forewing length average 36 mm; shape similar to others of the group except that 
the hind angle of the forewing is slightly less prominent in ib so that the angle is more 
rectangular. Upperside. Forewing, base and distal portion black, base slightly browner, 
white bar broader than in nominate race, 6 mm at costa expanding in a slight curve to the 
hind angle where the bar is 8 mm wide, the inner border more extended proximad and shaded 
with bluish grey, but reduced at the costa where there is a black line at end of cell. Hind 
wing, basal area and inner fold brownish black to anal angle where there is a small pale spot 
at inner margin. Outer border of wing blacker, extending from upper angle to above the 
tail. White band wider than in nominate race, 10 mm at costa, with a narrow white central 
line extending to base of vein 4 flanked proximally with greyish blue, less strongly blue 
distally, the white area ending rather at vein 3, giving way to the rufous-orange which extends 
to the hind margin and the tail; the ocellus in space 3 conspicuous, that above the tail less so, 
but the white dots at anal angle distinct though small. The main character is thus the width 



REVISIONAL NOTES ON AFRICAN CHARAXES 75 

and extent of the white bar on the hind wing. Underside. Very similar to that of the nominate 
race but fore wing white bar much broader throughout, the hind wing bar also wider. 

Female. Larger than the male, fore wing length 45 mm, shape as in nominate race. 
Upperside. Fore wing, basal dark area as in nominate subspecies, but distal dark border 
stronger, the arrow-shaped submarginal spots rufous-orange except that at the costa which is 
white to buffy. Hind wing, white band more defined distally, the outer border dark. The 
submarginal spots larger and rufous-orange; the ocellus conspicuous. Tail robust, rufous in 
colour but pale tipped, 10 mm long. Underside. Fore wing, very similar to that of the 
nominate race but the distal portion more tinged with buffy, the fine wavy dark lines 
commencing more abruptly. The submarginal spots buffy though rather obscured. Hind 
wing, white band more defined distally by the dark wavy lines commencing more abruptly 
on the buff-tinged ground. Submarginal spots in the form of ocelli more rufous but less edged 
with white, but large ocellus at hind margin and anal angle well marked. 

Holotype <$, Uganda: West Bwamba Valley, v. 1954 (van Someren) (BMNH). 

Paratypes. Uganda: Bwamba Valley, vii. 1942 (van Someren), 1 $ (allotype); 
Bwamba Valley, 1 <J (BMNH). 

Range. Uganda, Ruwenzori, Bwamba Valley. A transitional form occurs 
in the Epulu area of Zaire (PI. 5, fig. 34). 



Valla publius Staudinger 

Valla publius publius Staudinger 

(PI. 5, figs 38, 39, Text-fig. 5; Map 2) 

Palla publius Staudinger, 1892 : 267. 

Nearest in general fades to Palla ussheri (Butler), but exhibiting constant 
differences; the females of the two species being totally different. 

Male. Fore wing length 37-38 mm. Shape as in other species of Palla. Upperside. 
Fore wing, pattern very similar to that of P. ussheri, the white band slightly wider and slightly 
more curved on distal border. Hind wing, pattern also very similar but white portion of the 
band more extended, reaching vein 6, joining the rufous-orange rather abruptly and expanding 
rapidly at an angle and extending to the hind margin, including the whole of the tail. Underside. 
Fore wing, basal area and distal portion of wing darker than in other species, the fine dark lines 
on the latter, on a more brownish base, the margin of the wing darker. The same remarks apply 
to the hind wing. 

Female. Unlike other species, the female of publius resembles somewhat a larger paler 
edition of the male, having in general a similar pattern above. Fore wing length 45 mm. 
Upperside. Fore wing with an ochreous spot in the subapex in 5 and more obscured spots at 
the costa. Underside. Fore wing, paler than in the male, especially on the outer border 
where the fine wavy lines are on a brownish grey ground ; in addition, there are dark rays on 
the distal side of the white band, at veins ia-4, and dark lines on the submargin in the curve 
of the wing. The white bar very similar to that of upperside. Hind wing, white band very 
distinct, widest at the costa and tapering to above the anal angle; the ground colour of the 
dark wavy lines is buffy, but with dark patches in 4-5 and costa to 5. The marginal rufous 
ocelli, edged proximally in white, distinct, especially that in space 3 on the hind margin, showing 
up clearly on a clayish olive ground, proximally accentuated by a dark line. 

Range. Ivory Coast, Ghana, Sierra Leone to eastern Nigeria. 



76 V. G. L. van SOMEREN 

Form rectifascia Weymer 

Palla rectifascia Weymer, 1892 : 91. 

This form, in both sexes, only differs from the nominate race, in having on the upperside 
a straighter fore wing bar, but the white area of the hind wing bar extends further into the 
orange on the proximal side, reaching vein 5. However, in the female the white bar of the 
fore wing is broader throughout, the submargin with a row of tawny spots, that in the subapex, 
whitish. On the hind wing the white upper part of the band is broader and extends further on 
the proximal side, almost reaching the inner fold where it merges with the orange; the 
submarginal spots obscure at upper half, but more distinct towards the hind border where the 
black ocellus is prominent. Underside, very similar to that in the nominate race but dark 
patches on fore wing stronger, on a paler ground. Hind wing ground colour less buffy but 
the admarginal ocelli well marked. 



Form moderata Gaede 

(PL 6, figs 49, 50) 

Palla moderata Gaede, 19 16 : 71. 

The type of P. moderata Gaede cannot be located, and the description was not 
accompanied by a figure. A translation of the description is as follows. 

Closely related to P. ussheri Butler. The white band of the fore wing is slightly narrower; 
inwards, it has a slight bluish margin. The white band of the hind wing extends from the 
anterior margin to R 7 and then it becomes brownish. This brown part of the band is 
considerably narrower than in all Palla spp. and does not become broader towards the inner 
angle, but maintains an even width from the outer margin. The band is separated on the 
inner angle from the reddish yellow spot that extends from the inner margin to R 7 on the 
margin. In other species this spot is united to the band. An isolated spot is present in 
areas 4 and 5, the former exhibiting a white centre. The spot in area 4 in ussheri is nearly 
always united with the band, only in rare cases is it separated. The lower surface of the 
hind wing is identical with that of ussheri. This means that the differences between the 
two species is not very great, and it is not unlikely that some day transitional specimens 
will be found. However, it is also impossible to differentiate publius Stgr. and ussheri, 
by comparison of the upper surface while the lower surface differs slightly. In spite of that, 
the remarkable difference of the females confirm that they are separate species. 

The female of moderata is unknown. 

Type male, Dengdeng, N. Kamerun. 3/4/14. Coll. Dr. Milbraed. In addition, one male 
from Sierra Leone, in collection Staudinger. 

It will be noted that these two specimens assigned to moderata come from widely 
separated areas. A third specimen placed under moderata in the Rothschild 
collection (BMNH) has been sent to me by Mr Howarth. It was taken at Kapu- 
lumbo, Kasai, western Congo, also far distant from Cameroun. This specimen 
closely resembles P. publius on the upper side, more so than ussheri, and suggests 
that moderata Gaede is not a species but a variation of publius, occurring here and 
there within the range of that species. 

This specimen does not quite agree with the description of the type in so far as 
the orange band on the hind wing upperside is concerned, for it widens considerably 
in its lower portion above the anal angle. 



REVISIONAL NOTES ON AFRICAN CHARAXES 77 

Palla publius centralis subsp. n. 

(PI. 5, fig. 40; PI. 6, figs 41-45; Map 2) 

Male. Similar in size and shape to the nominate race, but differing in the following 
characters. Upperside. Fore wing, bar is narrower in the region of the costa and the cell 
end, being 3 mm or even less, but gradually expanding towards, but contracting slightly, 
at the hind margin, particularly on the outer edge so that there is not a large overlap of the 
fore wing bar over that of the hind wing. The inner border is less sharply defined due to some 
greyish blue scaling, and there is often a whitish mark in the cell end. Hind wing, the white 
area of the bar is widest at the costa, 5 mm decreasing to 2 mm where it encroaches within the 
commencement of the orange area which expands rapidly to reach the tail on the distal side 
and the anal angle on the proximal side, the inner side being almost straight. The ocellus 
on the hind margin is very distinct as are also the two white dots at the anal angle. The tail, 
though robust, is pointed with rounded tip. Underside. Fore wing, very similar to that of 
the nominate, the white lines crossing the cell are distinct though the basal area is not so dark. 
The white bar is similar to that of upperside. The fine lines in the postdiscal zone are more 
distinct on a less dark border. Hind wing, the white bar is narrower, especially at the costal 
end, and the bar is of almost even width and more curved; otherwise, the pattern follows that 
of the nominate race. 

Female. Here also, it is male-like. Length of fore wing 45 mm. Upperside. Differing 
from the female of the nominate race in having a narrower bar throughout. Forewing, postdiscal 
row of spots obscured or absent. Hind wing, the white area at the costa is narrower and 
less extended, but the orange area expands rapidly as a triangle to reach the hind margin 
including the tail and the anal angle; the ocellus is very distinct. Underside. This exhibits 
similar differences to those noted in the male, but on the hind wing the submarginal ocelli 
above the tail are not so distinct. 

Holotype $, Cameroun: Johann-Albrechts Hohe, Station Kamerun, 1896 
(L. Conradt) (BMNH). 

Paratypes. Cameroun: no further data, 1 $ (allotype); Bitye, 2000ft, Ja 
River (G. L. Bates), 1 <J; Mamfe, xi. 1956 (T. H. E. Jackson), 1 <J; Bitje, Ja River, 
2000 ft, dry season (G. L. Bates), 1 $. Zaire: no further data, 1 $. (All BMNH.) 

Range. Cameroun, Central African Republic, Zaire. 

Palla publius kigoma subsp. n. 

(PI. 6, figs 46-48; Map 2) 

Male. Differs from both the nominate race and p. centralis by the greater, more even width 
of the fore wing bar, which is almost parallel sided, especially on the undersurface. The white 
area of the bar on the hind wing is also wider as is also the orange portion of this band. On 
the underside, the pattern is strong. 

Holotype $, Tanzania: Kigoma, Kabogo, 28.xi.1961 (Japanese Primate 
Expedition) (BMNH). 

Paratype. Tanzania: holotype data (Nat. Mus. Nairobi, Kenya), 1 <$. 

SYSTEMATIC LIST 

Palla decius (Cramer) 

Palla decius decius (Cramer, 1777). Type-locality: Coast of Guinea. 

f. Sagittarius Rousseau-Decelle, 1934. Type-locality: Zaire, Kasai district. 
Range: Guinea, Sierra Leone, Congo (Brazzaville), N. Angola. 



7 8 V. G. L. van SOMEREN 

Valla ussheri (Butler) 

Palla ussheri ussheri (Butler, 1870). Type-locality: Gold Coast [Ghana]. 
f . ferruginea Schultze, 1914. Type-locality: Cameroun. 

Range: Sierra Leone to W. Nigeria, west of Cross River. 
$ f. dobelli Hall, 1919. Type-locality: Cameroun, Bitje, Ja River. 

Range: Cameroun, Central African Republic, Congo (Brazzaville). 
ussheri interposita Joicey & Talbot, 1925. Type-locality: Uganda, Mabira 
Forest. 

Range: Uganda, from west to east but mainly in central forests; Zambia. 

Palla violinitens (Crowley) 

Palla violinitens violinitens (Crowley, 1890). Type-locality: [Ghana] Accra. 
Range: Ghana to Central African Republic. 
violinitens cline coniger (Butler, 1896). Type-locality: Old Calabar (<£), 
Zaire ($). 

Range: Cameroun, Central African Republic, Congo (Brazzaville), Kasai 
and central Zaire. 
violinitens bwamba subsp. n. Type-locality: Uganda, Bwamba Valley. 

Range: Uganda, west of Ruwenzori, Bwamba Valley with variation in 
E. Zaire in the Epulu area. 



Palla publius Staudinger 

Palla publius publius Staudinger, 1892. Type-locality: Sierra Leone, 
f. rectifascia Weymer, 1892. Type-locality: Ghana. 

Range: Ivory Coast, Ghana, Sierra Leone to E. Nigeria, 
f. moderata Gaede, 1915. Type-locality: Cameroun. 

Range: Sierra Leone, Cameroun, Zaire. 
publius centralis subsp. n. Type-locality: Cameroun, Bitje. 

Range: Cameroun, Central African Republic and Zaire. 
publius kigoma subsp. n. Type-locality: Tanzania, Kigoma. 

Range: Tanzania. Only known from the Kigoma district north-east 
of Lake Tanganyika. 



THE EUXANTHE COMPLEX 

The genus Euxanthe was created by Hiibner in 1816 (1819), without a generic 
description. The type-species was cited as Papilio eurinome Cramer (1775-6). 

The genus was subdivided by Aurivillius (1898 : 220) when he created the sub- 
genus Hypomelaena (type-species Godartia trajanus Ward, 1871). The distinguishing 
characters are that in Euxanthe the fore wing cell is obtusely rounded and elongated 
at its distal end, the hind wing ceU is open, while in Hypomelaena the fore wing cell 
is almost triangular and ends abruptly and the hind wing cell is closed. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



79 



A great deal of reliance is placed on wing venation for distinguishing genera, 
and rightly so, but, in my opinion, the early stages are also of great importance. 
Wing shape is also of significance in 'grouping' and, although the shape of Charaxes 
and Euxanthe differ considerably, evidence is afforded by the ova, larvae and pupae 
which show that Euxanthe is allied to Charaxes, but forms a compact group. I 
therefore support the proposal of Rydon (1971 : 230) that a sub-family Euxanthinae 
be erected for the group. For comparative figures and descriptions of the early 
stages vide van Someren & van Someren (1926 : 354) and Rydon (1971). (See 
Text -figs 7-9.) 

Euxanthe {Euxanthe) eurinome (Cramer) 

Euxanthe (Euxanthe) eurinome eurinome (Cramer) 

(PI. 7, figs 51-54; Map 3) 

Papilio Eques Achivus eurinome Cramer, 1775 : 109. 
Papilio Festivus eurinome Cramer; Fabricius, 1793. 
Euxanthe eurinome (Cramer) Hiibner, 1816 : 39. 
Euxanthe eurinome (Cramer); Kirby, 1871 : 228 

For full references vide Stichel, 1939. 

Male. Fore wing length 42-44 m. Shape, wings rounded, costa and outer margin of wing 
curved, apex rounded, or very slightly pointed. Upperside. Fore wing, ground colour black 











8 

Figs 7-9. Early stages of (7) Charaxes, (8) Euxanthe, (9) Palla. (R. Smiles del. after van 
Someren.) 



8o V. G. L. van SOMEREN 

slightly browner at base and costa. Pattern consisting of four rows of spots, all slightly bluish 
green, the intensity of the colour depending on the direction of the light upon them; row i, 
mark towards end of cell small and elongate with a small dot at proximal end, an elongate 
ovoid mark in space 2, followed by two narrow elongate marks in space ib and a smaller mark 
in ia placed more proximad; row 2, elongate marks in spaces 3-6, that in 5 a narrow streak; 
row 3, with four smaller elongate spots; submarginal spots smaller and whitish, rounded, from 
subapex to space 3, the mark in 2 more angular and set in, two spots in ib. Hind wing, ground 
colour black, the basal area with a large bluish green patch occupying the cell and space below 
and bases of 5-6, the marks in the latter separated by black, so that the outer border is irregular ; 
postdiscal line with conspicuous row of ovoid bluish green spots, largest at costal end, smaller 
in 4—5, then larger in 2—3; the spots on the inner fold small; the marginal row of spots more 
rounded, the spots decreasing in size from upper angle, double spots at anal angle; edge with 
small white marks opposite each spot. Underside. Fore wing, ground colour dull black, 
more brownish along the costa to apex, and the outer margin to hind angle. Pattern of spots 
as upperside but paler, but cell and base of costa with whitish spots; the submarginal spots 
in spaces ib and 2 more angled and larger than above. Hind wing, ground colour dull black 
shading to brownish at base of costa and upper part of inner fold ; basal light area larger than 
above, the whitish extending into the inner fold from ia-ic; the postdiscal row of pale spots 
larger and more silvery white, and in addition a series of small white spots in the interspaces. 
Female. Fore wing length 45-51 mm; shape less rounded than in the male, the outer border 
less outwardly curved. Upperside. Fore wing, pattern of pale spots arranged as in the male, 
but all spots larger and white, with just the slightest tinge of green. The mark in the cell 
large and elongate, sometimes filling the whole of the cell, but usually with an irregular upper 
edge; the spot in space 2 large, and extending basad; the mark in 3 in the mid row large, rounded 
or ovoid, often extended basad; the submarginal spots larger than in the male. Hind wing, 
ground colour black, browner along the costa and along the distal border of the large white 
area in the disc of the wing which extends into the inner border to above the hind angle, the 
outer edge slightly dyslegnic, represented at the costa by an oval free spot, white in colour. 
The postdiscal row of more rounded spots, white and distinct and, unlike the male, there is a 
double row of white spots between the postdiscal row and the small white spots on the admargin. 
Underside. Fore wing ground colour, dull black, the pattern of white spots more or less as 
upperside, the cell mark often with a black spot in subbase. Hind wing ground colour dull 
brownish black, the border slightly blacker; the basal white area more rayed by blacker veins, 
but not extending so far down on the inner fold; the postdiscal row of white spots larger and 
more rounded than upperside, the intermediate submarginal white spots as upperside but 
larger and extending from costa to hind margin at the angle; the submarginal white spots 
larger and more elongate; the edge with a narrow white fringe opposite these spots. 

Range. The nominate race extends from Sierra Leone, Ivory Coast and Ghana 
to Nigeria; it also occurs on Fernando Po. There is reason to believe that nominate 
eurinome intergrades with the more eastern race ansellica in the region of Cameroun, 
the Central African Republic and Zaire. 



$ form johnsoni Howarth 

Euxanthe eurinome Cramer ^ form johnsoni Howarth, 1969 : 154, pi. 2 fig. 5. 

This extreme female form is distinguished by the enlargement and coalescing of all the 
white markings basad of the submarginal spots into one large area on the upper and underside 
of the fore wings. 



REVISIONAL NOTES ON AFRICAN CHARAXES 81 

Euxanthe (Euxanthe) eurinome ansellica (Butler) 
(PI. 7, figs 55-58; PI. 8, fig. 59-63; Maps 1, 3) 

Godavtia ansellica Butler, 1870 : 525. 

Euxanthe eurinome ansellica var. radiata van Someren & Rogers, 1927 : 65. 

Male. Fore wing length 41-45 mm. Shape as in the nominate race. Upperside. Fore 
wing, pattern and colour of bluish green spots very similar, the spot in space 3 set rather more 
distad, those in ib of more equal length; the spot in the cell larger and more irregular on upper 
border; the general ground colour similar to that of the nominate race. Hind wing ground 
colour as in eurinome eurinome, the general pattern very similar, the second spot in space 5 
of the discal row often missing. Basal patch often broken up by black and the whole patch 
is more limited on the proximal side ; the spots in the discal line and admargin as in the nominate 
race. Underside. Forewing, ground colour slightly more brownish than in nominate race; 
the pattern also very similar, but submarginal row of spots on the hind wing stronger and 
more developed. 

Female. Larger than the male, fore wing length 54-55 mm; shape slightly more convex on 
outer border. Upperside. Fore wing; this sex exhibits a marked departure from the pattern 
of the nominate race, the pale markings on the fore wing are smaller, thus the spots are more 
scattered, and all are pale bluish white. Hind wing, the basal patch is less solid and is separated 
into pale bluish white rays by the black-scaled veins; however, the submarginal and admarginal 
bluish white spots are more developed. Underside. Ground colour browner than in nominate 
race, the pale spotting, though larger than on upperside and pale bluish in colour, is similarly 
broken up. The colour of the abdomen is darker dorsally. 

Range. Northern Angola, S. Cameroun, Central African Republic, central 
and southern Zaire including Katanga; extending eastwards to west and central 
Uganda. 

Specimens from eastern Zaire have received the name /. burgeoni Le Cerf, 1923, 
the type-locality being Kindu. This form was placed under eurinome eurinome 
by Peters (1952), which is incorrect (PI. 8, fig. 64). 



Euxanthe {Euxanthe) eurinome celadon Le Cerf 
(PI. 8, fig. 65; Map 3) 
Euxanthe eurinome celadon Le Cerf, 1923 : 363. 

There has been some confusion in the past as to the exact locality of the race 
celadon Le Cerf. Specimens from the Mt Elgon area having been placed under it 
because of their small size, compared with ansellica. In order to clarify the position, 
I wrote to Dr Viette of the Museum National d'Histoire Naturelle, Paris, and asked 
him to kindly check on the locality cited on the label attached to the type-specimen. 
This label gives the locality as Gabun! Bryk (1939) gives the type locality as 'Congo' 
without exact location. Since the race ansellica Butler occurs to the immediate 
east of Gabun, extending east through the Congo to Uganda, it appears obvious 
that the Elgon insect cannot be placed to celadon. 

Unfortunately, I have no topotypical specimen of celedon on which to base a 
personal opinion as to its validity; I accordingly quote the original description 
given by Le Cerf. 



8z V. G. L. van SOMEREN 

Euxanthe eurinome Cr. s. sp. celadon nova. 

cJ.-Aire basale verte des ailes inferieures fortement reduite par l'envahissement du fond 
noir qui couvre largement les bords de la cellule et les nervures; la tache entre i b et la cellule 
n'atteint pas la base de la nervure 2; celle de l'intervalle 3 manque ou est rudimentaire ; 
celles des intervalles 6 et 7 sont en majeure partie obliterees par un semis noir. 

9- - Tous les dessins d'un bleu verdatre clair au lieu de blanc pur; bande terminale noiratre 
des ailes inferieures large, remplissant la base des nervures 2-3 ; bords de la cellule et nervures 
ecailles de noir; entre les nervures i b et 2, un large trait noir court sur le pli jusqu'a 1 
centimetre de la base de l'aile. 

Types : $ (H.T.), Gabon, ex M. Lebaudy, 1909. - 1 £, 1 <j>, Tchibanga, Gabon, 1908, ex 
G. Le Testu. - 1 $, 1 $, Landana, Congo portugais, 1882; 1 $, Thoumby, Congo beige, 1883, 
ex L. Petit < coll. E. Boullet, Coll. Museum National de Paris. 

Range. Gabon. 

Euxanthe {Euxanthe) eurinome elgonae subsp. n. 

(PI. 9, figs 66-73; Maps i, 3) 

Male. Fore wing length 40-42 mm, thus smaller than the nominate race and ansellica of 
the Congo and central Uganda. The chief characters of the race elgonae are its considerably 
smaller size compared with ansellica, but the pattern of the male on the upper side is more like 
that of nominate eurinome, although the underside ground colour is quite different. The female 
of elgonae is more like that of the nominate race regarding the solidity of the hind wing patch, 
which is white, but the fore wing spots are blue as in ansellica. In detail, the differences com- 
pared with ansellica are as follows. Upperside. Fore wing, cell mark is divided into two, 
one at subbase and a double spot beyond; though somewhat variable, the postdiscal and 
marginal spots are more uniform in size ; hind wing, the basal green patch is smaller and more 
consolidated distally, less broken up by dark rays. The postdiscal row of spots more rounded 
and uniform in size; the submarginal row of small spots better developed. Underside. Fore 
wing ground colour, brownish along the costa and the outer border. Hind wing, more 
brownish along the costa. 

Female. Fore wing length 45-50 mm. Upperside. Fore wing, pattern of pale spots 
bluish to whitish. The hind wing patch is white and more consolidated, its outer border 
strongly indented in spaces 2-3. Underside. Hind wing, ground colour very similar to that 
of ansellica, but pattern reflecting the differences noted on the upperside. 

Holotype $> Kenya: east Elgon area, Trans. Nzoia district (BMNH). 
Paratypes. Kenya: holotype data, 1 $ (allotype) (BMNH), 1^,1$; Mara 
District, Gori River forest, 2 <$. Uganda: W. Madi, Metu forest, 1 <J. 

Range. North-west Kenya to the east and south-east of Mt Elgon, extending 
north into the Suk country. Similar specimens have been taken in Uganda in the 
West Madi district at Metu. Its distribution coincides with that of its chief food 
plant, Mbambakofi, Afzelia africana Smith (Caesalpinaceae) on which it has been 
bred. 

Euxanthe {Euxanthe) eurinome birbirica Ungemach 

(Map 3) 
Euxanthe eurinome birbirica Ungemach, 1932 : 52. 

As I have no specimen of this subspecies I quote below the original description. 



REVISIONAL NOTES ON AFRICAN CHARAXES 83 

82. Euxanthe eurinome birbirica s.-sp. nova. 

La race abyssine de ce bel insecte se rapproche de la race congolaise ansellica Btlr. par 
la reduction de l'aire basale claire de l'aile posterieure et par l'irregularite de la serie de 
taches submarginales de l'aile anterieure. Mais les taches submarginales de l'aile post, 
sont de la taille de celle d'eurinome Cr. et presque rondes. Toutes les taches claires de l'aile 
ant. sont beaucoup plus reduites que dans les deux races connues; en particulier, la grande 
tache cellulaire se divise en deux petites taches, une triangulaire a la base, un autre allongee 
et bilobee au dela du milieu. La tache de l'intervalle i a est absente ou minuscule. 

Chez la femelle, les taches de l'aile ant. sont blanc bleute, la base de l'aile post, blanc pur 
et les taches discales vert tres pale. Mais surtout, le fond de l'aile post, est brun rouge au 
lieu de noir, presque de la meme teinte que le dessous du $. L'extremite de l'intervalle i b 
est meme brun clair. Les arceaux noirs de la bordure envoient un prolongement noir 
jusqu'aux taches discales. II n'y a que deux discales dans l'intervalle i c , dans les deux sex. 
Envergure de la $, 97 mm. - Type dans ma collection. (Youbdo, 13 nov. 26). 

J'ai capture de cette belle espece trois $$ et une 9 a. Youbdo; tous, sauf un 3, sur un 
arbre blesse dont la seve en fermentation attirait de nombreux insectes et surtout une grande 
quantite de Charaxes. 

Euxanthe (Euxanthe) crossleyi (Ward) 
Euxanthe (Euxanthe) crossleyi crossleyi (Ward) 

(PI. 10, figs 74-80; Map 3) 

Godartia crossleyi Ward, 1871 : 36. 
Euxanthe crossleyi (Ward) Kirby, 1877 : 740. 
Euxanthe crossleyi f. niepelti Bryk, 1939 : 632. 

Male. Fore wing length 45 mm; shape, costa curved, apex only slightly pointed, outer 
margin outwardly curved. Hind wing rounded, no anal projection. Upperside. Fore wing 
ground colour black; pale markings in four rows, all pale greenish cream, cell almost entirely 
filled by a large black mark but not reaching the base, sometimes with a black dot in mid- 
subcostal area, followed by a large rather triangular mark in space 2, followed by narrow 
elongate marks in ib-ia; long narrow marks in subbases of 4—6, mark in 3 at a slight angle to 
those above and set in a little; postdiscal row of spots smaller, in a row from 4-7, linear; 
submargin with full series of more rounded spots from ib to subapex in 7, the spot in 2 largest 
and set in a little, mark in ib double; base of fore wing costa with a pale line, sometimes tinged 
with rufous basally. Hind wing, ground colour black, disc of wing filled by a large greenish 
cream patch separated up into rays by black veins, the subcostal mark in 7 very long and 
extending distad to fuse with the postdiscal row of large ovoid pale spots, with marks below 
also fusing to a lesser degree; represented on the inner fold by a club-shaped whitish mark 
which is freckled with black scales. Submarginal line with a row of double half-moon spots 
divided by black; admargin with pale linear marks in interspaces. Underside. Fore wing 
ground colour black, slightly paler along the costa, apex and outer border. Pattern of pale 
spots arranged as upperside but paler. Hind wing, base of costa and 6 with a rufous patch with 
two white dots; rest of the wing taken up by the paler greenish cream enlarged patch as on 
upperside, divided up by black veins, the inner fold greenish cream divided longitudinally 
by black veins, not all black as upperside. 

Female. Larger than the male, fore wing length 58-60 mm. Shape, outer margin less 
outwardly curved than in the male. Upperside. Fore wing, basal part of wing taken up 
almost entirely by the pattern of creamy marks which are only slightly tinged greenish in the 
distal part of the wing. Black triangular area at base of cell, extending into bases of spaces 
ib-ia. Cell itself entirely creamy; the discal zone of the wing with an enlarged creamy area 
divided up by black veins; the postdiscal series of spots from 4 to subcosta in 7, much larger 



84 V. G. L. van SOMEREN 

than in the male, and creamy in colour; the submarginal spots arranged as in the male but 
larger and more angular. Base of costa with a creamy line. Hind wing, extreme base black, 
bulk of wing filled with the creamy rayed pattern, which is more extensive than in the male, 
the ends of the rays more fused with the postdiscal rounded creamy spots. The inner fold is 
creamy, divided by black veins; the submarginal and admarginal creamy spots arranged as 
in the male but larger. Underside. Fore wing, ground colour towards the apex and outer 
border greyish, darkening toward the hind angle; the black patch at lower base of cell and 
base of ib strong. The bulk of the wing filled by an enlarged pattern of creamy marks arranged 
as on upperside; the postdiscal spots more or less fusing with the discal spots. Hind wing, 
ground colour more greyish, the whole area taken up by the extended pattern of creamy rays 
which fill it from the base to the submarginal zone, where the row of pale spots is strongly repre- 
sented as are the admarginal triangular spots. The extreme base of the wing is shaded with 
rufous. 

Range. The nominate race occurs in the Cameroun, Zaire, the Central African 
Republic and Mwinilunga, Zambia. 

Euxanthe (Euxanthe) crossleyi magnified Rebel 
(PI. 12, figs 90-92; Maps 1, 3) 

Euxanthe crossleyi magnified Rebel, 1914 : 252. 
Euxanthe crossleyi intermedia Joicey & Talbot, 1921 : 75. 

Male. Fore wing length 40-42 mm; shape as in the nominate race. Upperside. Fore 
wing differs from the more western crossleyi crossleyi by its paler coloured pattern, the greenish 
tinge to the creamy marks being mostly on the distal half of the wing. The general pattern 
very similar to that in the nominate race but more extended; ground colour black. Base of 
fore wing costa with a marked creamy stripe up to mid point ; cell filled entirely with creamy 
yellow except for a streak at lower base adjoining the black spot at the base of the wing which 
fills the bases of spaces ib— 3, as in the nominate race, but more extended, that in 3 fusing with 
the spot in the discal row ; the marks in the discal row larger and more extended ; the four post- 
discal spots larger as also are the submarginal spots, of which that in 2 is slightly set in and larger, 
that in ib double. Hind wing, pattern as in the nominate race but creamy, the rays of the discal 
area more even and elongate, the fusion with the spots in the postdiscal row more complete; 
the spots in the submarginal line larger and creamy, as also are the admarginal ones. The 
inner fold of the wing creamy buff, divided by black veins. Underside. Fore wing ground 
colour shading to greyish on the apex and outer border; the heavy black mark at the base 
strong. The general pattern follows that of upperside but is paler in colour. The hind wing 
pattern is essentially that of upperside, divided by black veins. The ground colour on the 
border is more greyish; the submarginal and admarginal spots as upperside but paler. 

Female. Larger than the male, but with similar pattern and all spots paler. 

Range. Eastern Zaire, mainly in the Irumu-Beni-Ituri areas. It is of interest 
to note that this pale race occurs between the darker western and the strongly- 
marked race of Uganda. 

Euxanthe {Euxanthe) crossleyi ansorgei Rothschild & Jordan. 
(PL 11, figs 81-86; PI. 12, figs 87-93; Maps 1, 3) 

Euxanthe crossleyi ansorgei Rothschild & Jordan, 1903 : 333. 
Euxanthe crossleyi ansorgei i. babbingtoni Stoneham, 1943 : 46. 

Hitherto, all representatives of the species crossleyi from Uganda and Kenya 



REVISIONAL NOTES ON AFRICAN CHARAXES 85 

have been considered to be a homogeneous entity. After intensive study they 
may be divided into three groups. 

Group A. The type of ansorgei came from the Nandi forest. Both sexes are 
characterised by the smallness of the spots in the fore wing and the restricted 
hind wing patch, and the large postdiscal spots, but small admarginal ones. The 
streak on the hind margin in space ia is short. Topotypical examples: PI. 11, 
fig. 86; PL 12, fig. 88. 

Group B. The aggregate from the low-lying humid swamp forests of Katera, 
on the western shores of Lake Victoria, in the Masaka district, differ by having a 
large spot in the fore wing cell, and larger marks in the discal line, but smaller spots 
in the upper postdiscal row. The streak on the inner margin in ia is much longer. 
On the hind wing, the discal patch is larger, being extended distally, and also on the 
inner fold; the submarginal spots larger. These general differences apply equally 
to the females. Specimens from central Uganda, Mawakota area, come within 
this group (PL 11, figs 81-83). (See Map 1.) 

Examples from the Kigoma district to the north-east of Lake Tanganyika in 
Tanzania, of which I have insufficient material, approach the Katera aggregate 
(B) but differ appreciably; the fore wing spots are strongly greenish (PL n, fig. 84). 

Group C. Moving further west to the forests of Kayonza in the Kigezi district 
of Uganda, we note that males differ from either of the above groups; the base 
of the fore wing costa is rufous, extending as a streak at the base of the cell; the 
mark within the cell is larger than in those in group A, and more broken up and 
irregular on the upper edge than in those in group B. The discal row of spots 
as in group B, thus larger than in group A; the postdiscal spots as in B but limited 
to three spots. The hind wing patch intermediate between that of A and B, but 
the postdiscal row of spots large as in A. The submarginal spots are small, but 
the admarginals are large and reach the margin. Females correspond in a like 
manner (PL 11, fig. 85; PL 12, fig. 87). (See Map r.) 

These aggregates therefore exhibit characters tending toward the two well 
marked races magnified and ansorgei, yet are sufficiently distinct from either and 
between themselves, as almost to warrant recognition as subspecies. 

Euxanthe (Euxanthe) crossleyi claudiae Rousseau-Decelle 

(Map 3) 
Euxanthe crossleyi claudiae Rousseau-Decelle, 1934 : 228. 
No specimens available for a personal assessment. 

Euxanthe {Euxanthe) wakefieldi (Ward) 
(PL 13, figs 94-100, Text-fig. 1; Map 3) 

Godartia wakefieldi Ward, 1873 : 152. 
Euxanthe wakefieldi (Ward) Kirby, 1877 : 740. 
Euxanthe wakefieldi f. rubiginea Le Cerf, 1923 : 363. 

Male. Fore wing length 40-43 mm. Shape, fore wing costa strongly curved, apex bluntly 
pointed, outer margin of wing outwardly curved. Hing wing rounded, margin slightly 



86 V. G. L. van SOMEREN 

undulating. Upperside. Fore wing, ground colour black, very slightly brownish at base 
and along the costa. Pattern of greenish spots strong but somewhat restricted ; a large ovoid 
spot toward the end of the cell and occasionally a spot at upper subbase ; discal row of greenish 
spots of varying shape as follows : an ovoid subcostal spot followed by a small streak in space 
5, a longer streak in 4, the marks in 3-2 much larger, the proximal ends extended towards 
the cell, the distal ends incised or oblique, the double mark in ib projecting distally towards 
the hind angle, a small linear mark in ia. All these marks are satin-green, but appear satin- 
white in oblique view. The postdiscal row of marks limited to three large ones in the subapex 
in 4-6 with an occasional small dot in subcosta; submarginal spots variable, spots in 2-3 
always present, those in 5-6 smaller, often obscured, the mark in ib usually absent. Hind 
wing, ground colour black, slightly brownish at base and along inner fold. Discal green-white 
patch, sometimes satin-white according to light, somewhat restricted, divided on distal border 
by black veins. Postdiscal row of rounded spots complete, decreasing in size from subcosta 
to the double spot in the hind angle; submarginal spots mainly on angle, but obscured beyond; 
admarginal spots small, limited in number or obscured. An occasional variant has all the 
spots very pale, with little or no greenish tinge, so that the pattern appears white. Underside. 
Fore wing, ground colour blackish in areas ia-3, shading to dull rufescent brown on the costa, 
apex and outer border. The pattern of marks as upperside, paler, and slightly larger; the 
spot at the base of the cell strongly marked. Hind wing, ground colour rufescent brown, 
the pattern as on upperside but spots larger, the patch in the disc of the wing extended into 
the inner fold; an additional mark on the costa, and two white dots at its base. The postdiscal 
and submarginal spots more complete and distinct than on upperside and whiter in colour. 

Female. Larger than the male, fore wing length 50-52 mm. Shape of fore wing more 
elongate than that of male, the outer margin slightly incurved at 2-3. Upperside. Fore 
wing ground colour jet black; pattern of spots in the disc of the wing similar to that of the male, 
but all marks larger and pure white, occasionally with a slight bluish tinge to cell mark and 
subapical spots which are large in spaces 4-6, with an occasional dot in 7. Submarginal spots 
limited to 2-3. Hind wing, costal area and outer border jet black, the disc filled by a somewhat 
rounded white area which extends into the inner fold, the upper part of area clear-cut, the 
lower border more dyslegnic due to the intrusion of blackish scaling; the white spots in the 
discal and submarginal rows complete and clear, those on the admargin small but distinct. 
Underside. Fore wing, ground colour mostly dull blackish brown along the costa and apical 
region. Pattern as upperside but slightly enlarged; the cell with a basal white spot. Hind 
wing ground colour brownish, the large discal patch clear-cut, but divided by black veins; 
postdiscal and submarginal rows of white spots complete. The edge of the wing with slight 
white fringe between veins. 

The chief food plant is Mbambakofi, Afzelia cuanzensis Welw. (Caesalpinaceae) . 
For an account of the early stages vide van Someren (1935 : 172). 

Range. Coastal forests of Kenya and Tanzania, including the islands of Pemba 
and Zanzibar, extending to Mozambique as far south as Delagoa Bay and inland 
as far as Mbala (formerly Abercorn) , Zambia and has been seen by Mr F. Schofield 
in the lower Luangwa Valley (teste M. N. Mitchell, pers. comm. 1973). 



Euxanthe (Euxanthe) madagascariensis (Lucas) 
(PI. 13, fig. 101; PI. 14, fig. 102; Map 3) 

Godartia madagascariensis Lucas, 1842 : 299. 

Anthora amakosa Doubleday, in Westwood, Doubleday & Hewitson, 1850 : 283. 

Euxanthe madagascariensis (Lucas) Kirby, 1871 : 228. 



REVISIONAL NOTES ON AFRICAN CHARAXES 87 

Most closely allied to the continental species Euxanthe wakefieldi (Ward) of 
eastern Africa, but pattern of light greenish spots more limited. 

Male. Fore wing length 40-42 mm; shape similar to that of other Euxanthe species but 
apex pointed. Upperside. Fore wing, ground colour black, pattern as follows: two rectangular 
greenish spots in distal half of cell, followed by a broad diagonal band of elongate marks in 
the discal line, extending from the subcosta towards, but not reaching, the hind angle, the 
spot in space ib the smallest. The submarginal row of small greenish spots of diminishing 
size extends from the subcosta to ib. Hind wing, ground colour black; pattern limited to a 
conspicuous greenish white area occupying the distal half of the cell and the bases of cellules 
above, the patch appearing white in oblique light. Postdiscal row of large greenish spots 
decreasing in size from subcosta to 3. Underside. Fore wing, ground colour brownish, darker 
in hinder portion. Pattern as upperside, but spots larger. Hind wing pattern much as upper- 
side but with a few additional white spots at base and on the inner fold ; postdiscal spots arranged 
as upperside but larger and extending towards the inner margin; submarginal spots complete, 
but admarginal spots limited to two or three in the mid-margin. 

Female. Larger than the male and slightly different in shape, the outer margin of the 
fore wing less curved outwardly. Upperside. Pattern more or less as in the male, but all 
spots white. The broad white discal area on the hind wing very large, extending from near the 
base to well beyond the middle and covering the bases of cellules 2-6. Postdiscal spots and 
those on submargin as in the male but all are white. A cinnamon-brown area at the anal 
angle. Underside. Fore wing base black, shading to chestnut on the outer border and apex, 
the brown colour extending along the costa. White markings as upperside, but enlarged; 
the spots on the submargin ringed in black, especially distally. Hind wing, ground colour 
on costa and border chestnut-brown; two white spots at base of costa; discal white patch as on 
upperside but outer margin more irregular; postdiscal white spots as above; an additional 
row of small white spots present in the submarginal line and larger spots on the admargin 
in the interspaces. 

Range. Confined to the island of Madagascar. 

Euxanthe (Hypomelaena) trajanus (Ward) 

Euxanthe (Hypomelaena) trajanus trajanus (Ward) 

(PI. 14, figs 103, 104) 

Godartia trajanus Ward, 187 1 : 36. 

Euxanthe trajanus (Ward) Kirby, 1877 : 740. 

Euxanthe schatzi Staudinger, in Staudinger & Schatz 1885 : pi. 48. 

Male. Fore wing length 45-46 mm. Shape, costa curved, apex rounded, outer margin 
outwardly curved; hind wing rounded. Because of the rounded apex, the fore wing appears 
more elongate than in other species of Euxanthe. Upperside. Fore wing base, the cell and 
base of space ib conspicously rufous-chestnut. Distal portion of wing black, with three rows 
of light marks, the discal row pale yellowish white, commencing with a large mark at end of 
cell, the distal end cut out to accommodate the oval mark at the base of 4, an angular mark 
fills the base of 3, followed by an elongate mark in 2, extending distad, a smaller triangular 
mark in ib with a smaller elongate mark below ; ia with a long white streak extending from base 
to just beyond the mid-point. Postdiscal series of large white spots, the three upper ones ovoid 
and large, the three lower more rounded, sometimes double at the hind angle, the lower spot 
very small; the subapex has two or three white spots diminishing in size, but occasionally 
present in 4 as a mere dot. Hind wing, ground colour black, the disc of the wing with a some- 
what restricted greyish white patch, clearly defined on its upper border, but dyslegnic on the 
outer, and more so on lower border toward the inner fold. The wing is otherwise immaculate 



88 V. G. L. van SOMEREN 

except for a series of small white dots on the admargin. The upperside of the abdomen is black. 
Underside. Fore wing, ground colour, matt black, base rufous-chestnut as on upperside; 
pattern of spots similar to that of upperside but subapical spots absent. Hind wing, ground 
colour matt black with slight brownish tinge, immaculate except for a minute white dot on the 
costa and mid point. 

Female. Upperside. Fore wing, somewhat similar in pattern to that of male, but mark 
in cell reduced to one small spot at subcosta and a spot in the lower apical region connecting 
up with the larger discal spot in the white band, which is very similar to that of the male but 
the mark at base of 4 is absent; the postdiscal row of spots like that of the male but spots larger; 
subapical spots limited to two. Hind wing, pattern mostly taken up by the large white area 
which extends from the inner fold, bordered distally by the black border which is slightly 
brownish at the hind angle. Admarginal row of white spots most marked in the curve of the 
wing. Underside. Fore wing, ground colour as on upperside, but black less intense and 
inclining to brownish at apex and on outer border. Base, chestnut area as on upperside; the 
discal white band as on upperside, but mark in cell increased in size costad, with an additional 
white spot beyond. Hind margin of wing with a short white streak. Hind wing, white area 
as on upperside but rayed by black veins and black lines in interspaces which run out to the 
margin, the surrounding ground colour brownish with black rays. Submarginal and admarginal 
spots as on upperside. 

Range. The nominate race is restricted to Cameroun and the countries bordering 
on the west of the Congo River. 



Euxanthe (Hypomelaena) trajanus vansomereni Poulton 
(PI. 14 figs 105-107; PI. 15, figs 108, 109) 

Euxanthe trajanus vansomereni Poulton, in Eltringham, Poulton, Riley & Talbot, 1929 : 476. 

Male. Fore wing length 47-48 mm ; shape, costa curved, apex blunt, outer margin outwardly 
curved; hind wing rounded. Upperside. Fore wing, general colour and pattern very similar 
to that of the nominate race from Cameroun, but rufous area of cell and basal area of space 2 
more extended distad, reducing the width of the yellow bar at end of cell and the other 
component yellow marks slightly reduced; the postdiscal white spots slightly larger; the sub- 
marginal spots as in the nominate race. The pale streak along the hind margin slightly longer. 
Hind wing, bluish grey discal area more extended proximally and distally and also towards 
the hind border, thus reducing the width of the black border of the wing where the submarginal 
white, double spots are larger and more distinct. Underside. Fore wing, ground colour 
not so black, more brownish; the pattern as upperside. Hind wing, ground colour more 
brownish so that the black rays show up more distinctly. The white dot at costa absent. 

Female. Fore wing length 60 mm. Shape as in nominate race. Upperside. Fore wing, 
pattern exhibits the same characters as noted for the male, the chestnut area in the cell is more 
developed, but the white band reduced in width, especially in space 3; the pale streak along 
the hind margin is narrower and tapering at both ends. The postdiscal white spots are more 
elongate, those toward the border in ib-3 smaller. Hind wing, discal patch creamy not white, 
more restricted on distal border which is straighter, not curved and does not extend so far 
down the inner fold; the black border is thus wider, and the two rows of white spots larger, 
especially those on the submargin. Underside. Fore wing, less black than in nominate race; 
between the white bars, more brownish. The basal chestnut more extended, but the discal 
white bar narrower; the postdiscal white spots more elongate. There is a double row of white 
spots in the apex. Hind wing, ground colour as in nominate race, black rays equally distinct 
and the distal border of the white patch straighter, as upperside; the two rows of white spots 
in the border larger and more developed, especially those on the admargin. 



REVISIONAL NOTES ON AFRICAN CHARAXES 89 

Range. Uganda, mostly in the central forests, extending east to the Mabira 
Forest. Specimens from Beni-Irumu, eastern Zaire, appear to be transitional, but 
the material available to me is poor (PI. 14, fig. 105). 

Euxanthe (Hypomelaena) trajanus gabonicus Le Cerf 
(PL 15, figs no, in) 

Euxanthe trajanus gabonicus Le Cerf, 1923 : 362. 

Euxanthe trajanus gabonicus i. depuncta Le Cerf, 1923 : 362. 

This subspecies is considered to be a synonym of nominate trajanus by the BMNH 
but I have insufficient material on which to base a personal opinion. 

Range. Gabon, Ogowe River. 

Euxanthe (Hypomelaena) trajanus antonius Rousseau-Decelle 

(PL 15, fig. 112) 

Euxanthe trajanus antonius Rousseau-Decelle, 1930 : 43. 

Through the kindness of Major Grahame, I am able to give a figure of a paratype 
specimen, ex coll. Rousseau-Decelle. It will be noted that the fore wing pattern 
is nearest to that of examples from the eastern Congo at Beni-Irumu, the chief 
differences being the more limited extent of the chestnut area in the cell, and the 
greater width of the creamy mark beyond; in these respects, the Katanga insect 
resembles the nominate trajanus. The hind wing greyish patch is more rounded, 
less angled on the distal side. 

Range. Southern Zaire (Katanga, Kafakumba). 

Euxanthe (Hypomelaena) trajanus nigeriae subsp. n. 



k ynypomeiaena) trajanus nigeriae sud 

(PL 15, figs 113, 114; PL 16, figs 115, 116) 
rth 4.3-44 mm: shape as in other races of traianu 



Male. Fore wing length 43-44 mm; shape as in other races of trajanus. Upperside. Fore 
wing, the chestnut area in the cell extends slightly into space ib; the creamy patch beyond 
in the cell end is separated from the chestnut by a black triangle, the rest of the creamy marks 
restricted in size; the whitish streak on the inner margin well developed; the white spots in 
the postdiscal line, bold; the subapical white spots, four in number, the lowest a dot. Hind 
wing, discal greyish area very i - estricted, rounded on outer border and merging with the inner 
fold ; the black border is thus wide ; the submarginal white dots small but distinct. For arrange- 
ment of pattern on underside vide PI. 15, figs 113, 114. 

Female. Fore wing length 52 mm. Upperside. Fore wing, chestnut area in cell limited 
to basal half; costal white spot small, that at lower angle elongate. The discal white area 
more solid, the lower edge straighter; the white spots beyond, as in the male, the lower three 
comparatively large compared with those above. The white streak on the hind margin strongly 
developed. Hind wing, basal area large, white in colour with only a slight tinge of rufous at 
hind angle, outer border very rounded; black border of wing thus reduced, but white sub- 
marginal spots distinct in upper half; admarginal spots punctiform but distinct. 

For underside vide PI. 16, figs 115, 116. 



go V. G. L. van SOMEREN 

Range. Nigeria. 

Holotype <$, Nigeria: Ikom, Ogolo Prov. iii.1956 (Jackson) (BMNH). 
Paratype. Nigeria: Ikom, Ogolo Prov., xi.1955 (Jackson), 1 $ (allotype) 
(BMNH) 

Euxanthe (Hypomelaena) tiberius Grose-Smith 

Euxanthe {Hypomelaena) tiberius tiberius Grose-Smith 

(PI. 16, figs 117-122, Text-fig. 2) 

Euxanthe tiberius Grose-Smith, 1889 : 129. 
Euxanthe tiberius f. tiberiella Strand, 1911 : 120. 

Male. Fore wing length 45-50 mm. Shape, costa curved, apex blunt, outer margin 
outwardly curved. Hind wing rounded, edge slightly undulating. Upperside. Fore wing, 
ground colour jet black, base with a large bright rufous-chestnut patch, filling most of the cell 
and the bases of spaces ia, ib and extending to the base of the costa. Pattern of greenish 
white spots in three rows, the discal row large, the spot at end of cell triangular, followed by a 
larger inverted triangle at base of 4, with a smaller triangular mark at base of 3, then a large 
elongate spot in 2 extending distad, followed by a horseshoe-shaped mark in ib; postdiscal 
row of more ovoid marks extend from costa to 3, more or less in line, then continued down the 
border as smaller spots; apex with ovoid spots decreasing in size to 4. Hind wing, ground 
colour uniform black, very slightly brownish at base of costa. The only pattern is a series of 
greenish white spots in the upper submargin, somewhat variable in number, those in 7-6 
being the largest and most constant; the admargin with a series of very small double dots, 
usually triangular in shape, often fading out at the upper angle. Underside. Fore wing, 
ground colour black in the discal area but browner in the apex and along the outer border. 
Base with the chestnut area as on upperside, the brown extending along the base of the costa; 
white dots present on base of costa and base of cell ; pattern of pale greenish white spots as on 
upperside. Hind wing, darker brownish with black rays along the veins and mid cellules. 
Triangular white dots present at base of wing and costa; spots on submargin and admargin 
as on upperside or limited to one at upper angle. 

Occasionally a dwarf specimen of either sex is taken in which the markings appear large in 
relation to the areas of the wings. 

Female. Fore wing length 50-53 mm. Shape somewhat like that of the male, but outer 
margin less outwardly curved. Upperside. Fore wing, ground colour black, with basal 
chestnut strong and well denned; the series of spots in the discal line pale greenish white 
arranged as in the male, somewhat variable in size, especially those at and beyond the end of the 
cell; the spots in the postdiscal line more whitish but also slightly variable. The spots in the 
submarginal line white and rather rounded, those in spaces 4 and 5 sometimes vestigial. Margin 
of wing with small white linear marks on the edge. Hind wing, a large white somewhat rounded 
area very slightly tinged with greenish, especially in the cell, the white area extending onto 
the inner fold but its upper border, which is rounded, starts just short of the dark base of the 
wing and cell, and does not reach the subcosta, its outer border more irregular, invaded by 
black along the veins. The border is widely black, with a row of white rounded spots in the 
submarginal line, that in 6 large and ovoid, the spots decreasing in size toward the hind angle; 
the admargin with a series of double angular white spots with a trace of smaller white lines 
nearer the edge which has an interrupted white fringe. Underside. Fore wing, ground colour 
black in the discal zone, shading to dull rufescent brown along the costa and the outer border. 
Base of wing with the rufous-chestnut patch as on upperside. Pattern of pale greenish white 
spots on upperside, those at the end of the cell slightly increased in size. Hind wing, ground 
colour rufescent brown, slightly darker on the border; discal whitish patch more clearly defined 
on border, invaded by dark veins and intermediary black lines which stop short of the edge of 



REVISIONAL NOTES ON AFRICAN CHARAXES 91 

the patch. Submarginal white spots as on upperside, those on the admargin outlined in black, 
the intermediary white dots more in evidence. 

For an account of early stages vide van Someren & Rogers (1928; 1932). 

Range. Coastal forests of Kenya (Rabai Hills, Shimba Hills, Marima Hill); 
also recorded from the lower forests of the Usambara Range at Amani, Tanzania. 



BIOLOGICAL NOTES 

By D. G. Sevastopulo 
(PI. 17, figs 129-136) 

Ovum. Spherical, the top fluted and only slightly flattened. Pale creamy when first laid, 
turning to a pale pinkish tan within 24 hours, finally becoming fairly dark brown all over. 
Deposited singly on the upper surface of a leaf of the foodplant. Laid 25.viii.64. Hatched 
31.viii.64. 

Larva. 1st instar. Head dark brown, rugose, the upper and lateral horns about the same 
length, slightly dentate, the lateral horns upcurved, the upper slightly divergent, incurved 
apically, the tips white. Body golden-brown, under a lens with a subdorsal, lateral and 
sublateral series of minute white papillae emitting colourless setae. Anal processes long, 
slender, curved, blackish tipped with white. Moulted 4.ix.64. 

2nd instar. Head dark brown with two paler transverse lines, the lateral horns both longer 
and stouter than the upper, two short spines between the upper, and a single spine between 
the upper and lateral and another below the lateral, all horns dentate and tipped with white. 
Body olive, very minutely papillated with white, the sublateral area tinged with crimson. 
Venter, legs and prolegs dark crimson. Anal processes as before. Moulted g.ix.64- 

3rd instar. Very similar to preceding, the two pale transverse lines on the head more 
noticeable and the horns longer, both actually and relatively. A fine white sublateral line 
above the crimson suffusion. Later in the instar a subdorsal white dot appears on the sixth 
somite. Body noticeably tapered from the ninth somite caudad. Moulted 14.ix.64. 

4th instar. Head whitish, two blackish transverse dentate bands, one just below the vertex, 
the other about central; upper horns slender, straight, slightly divergent, black at the base 
shading into greyish, the spines between black; lateral horns long and slender, horizontal, 
the tip upturned, colour white with the base and apex black, the spine between the upper and 
lateral horns white. Body much as in previous instar, the white subdorsal dots on the sixth 
somite ringed with black. Eighth somite with a smaller and less distinct white subdorsal dot. 
Anal processes no larger than before, and so proportionately smaller. After twenty-four 
hours the markings on the sixth somite become a whitish diamond, finely rimmed with black 
and containing two dark green dots. Moulted 20.ix.64. 

5th (final) instar. Head whitish, deeply punctate, each cheek with a diffuse olive spot divided 
vertically, a zig-zag olive transverse line on the level of the clypeus, which is filled in with pale 
green, and with two black spots set obliquely below it; horns similar in shape to previous instar, 
the lateral tipped with black and black behind, the upper black basally in front, the spines 
between the upper pair black, the jaws and posterior aspect of the head black. Body green, 
minutely shagreened with blue-white specks. Dorsal marks on the sixth and eighth somites 
diamond-shaped, pinkish white and studded with opalescent dots, edged with black and contain- 
ing two black spots, the whole distinctly raised above the general body level. Anal processes 
pinkish buff, short and stout, slightly incurved and joined by a transverse pinkish buff band. 
A pinkish buff sublateral line studded from second to ninth somites with fleshy whitish points. 
Legs green. Prolegs purplish. Venter green, shading into deep purple laterally. Pupated 
30.ix.64. 



92 V. G. L. van SOMEREN 

Pupa. Suspended by the cremaster. Mainly shining, rather translucent, dark green, 
marked with opaque bluish white as follows : a mark on the eye, a basal and submedian stripe 
across the wing joined by a connecting bar, and an irregular stripe along the outer margin, 
a broad transverse band across the third and fourth abdominal somites, containing a green, 
heart-shaped, dorsal mark on the third. Spiracles black ringed with white. Shape with the 
head slightly indented frontally, the thorax slightly keeled, wings laterally expanded across the 
tornus, abdominal somites rising to a transverse ridge on the third somite, which has a subdorsal 
prominence, and then tapering to the cremaster. Cremaster slender, green, with paired rounded 
projections on the dorsal aspect and a double projection ventrad. A male emerged n.x.64. 

Foodplant. Deinbollia sp. (Sapindaceae). 

Described from a larva reared from an ovum laid by a female caught in the 
Makadara Forest. 

Euxanthe (Hypomelaena) tiberius meruensis van Someren 
(PI. 17, figs 123-128) 
Euxanthe tiberius meruensis van Someren, 1935 : 172. 

Male. Fore wing length 44-46 mm. Shape as in the nominate race. Upperside. Fore 
wing, general pattern similar to that of t. tiberius, the rufous patch brighter and more extended, 
with a small yellowish dot beyond. Discal and postdiscal rows of spots, though similar in 
form to those of nominate tiberius, are all ochre-yellow. Submarginal spots white. Hind wing, 
ground colour slightly more brownish tinged, especially at base. Submarginal and marginal 
spots as in the nominate race. Underside. Fore wing, rufous patch brighter; distal portion 
of wing along the costa and apex brighter rufescent brown, with distinct black rays along the 
veins and intermediate spaces, the latter joining the submarginal subapical white spots with 
the ochreous ones. The discal spots formed as on upperside, all ochreous in colour on a black 
ground. A small white spot in margin. Hind wing, ground colour more rufescent brown; 
black rays slightly more distinct; submarginal and admarginal spots white with black surrounds. 

Female. Fore wing length 50-53 mm. Upperside. Fore wing, ground colour less intense 
black, more brownish black. The brownish rufescent area brighter and more extended. The 
discal row of spots narrower in spaces 2-3, all ochreous in colour. The postdiscal are also 
ochreous; the submarginal spots white. There is also an ochreous streak in the hind margin 
at about midpoint. Hind wing, ground colour less intense black, more tinged with brownish 
in the mid zone; the discal ochreous marking is narrower, being straighter on the hind border 
and not extending so much into the inner fold, the upper and outer borders straighter. The 
admarginal double spots white. Underside. Forewing ground colour in the apical half more 
rufescent and darker zone in the discal line not so black. The rufous patch at the base brighter. 
The spots in the discal zone ochreous; those on the submargin white. The streak on the hind 
margin similar to that of upperside, but larger. Hind wing, ground colour more rufescent 
brown, the discal patch restricted as on upperside, pale ochreous in colour. The black rays 
and veins distinct. Postdiscal spots as on upperside, ochreous in colour. Admarginal spots 
white with black surrounds. Edge very narrowly white in interspaces. 

Range. This very distinct race occurs in the forests on the lower slopes of 
eastern Mt Kenya, but is more plentiful in the lower Meru forest. It is also found 
in the forests in the Meru Game Park. 

SYSTEMATIC LIST 

Euxanthe (Euxanthe) eurinome (Cramer) 

Euxanthe {Euxanthe) eurinome eurinome (Cramer, 1775). Type-locality: 'Indies Orient' 
(patria falsa). Africa Occidens. 



REVISIONAL NOTES ON AFRICAN CHARAXES 93 

Range: Sierra Leone, Ivory Coast, Ghana, Nigeria, Fernando Po,? 
Cameroun. 
eurinome ansellica (Butler, 1870). Type-locality: Angola, Kinsembo. 
f. burgeoni Le Cerf, 1925. Type-locality: E. Zaire, Kindu. 
var. radiata van Someren & Rogers, 1927. Type-locality: Central 
Uganda. 

Range: Angola, Central African Republic, central and eastern 
Zaire, west and central Uganda. 
eurinome celadon Le Cerf, 1923. Type locality: Gabon. 

Range: Gabon. 
eurinome elgonae subsp. n. Type-locality: Mt Elgon. 

Range: N.E. Kenya in the Elgon area, Uganda, W. Nile district, 
Metu. 
eurinome birbirica Ungemach, 1932. Type-locality: Ethiopia, S.E. 
Youbdo. 

Range: S.E. Ethiopia, Youbdo district. 

Euxanthe (Euxanthe) crossleyi (Ward) 

Euxanthe [Euxanthe) crossleyi crossleyi (Ward, 1871). Type-locality: Cameroun. 
f. niepelti Bryk, 1939. Type-locality: Bipindi. 

Range: Cameroun, Central African Republic, Zaire, Gabon, Zambia. 
crossleyi ansorgei Rothschild & Jordan, 1903. Type-locality: N.W. Kenya, 
f. babbingtoni Stoneham, 1943. Type-locality: N.W. Kenya. 
Range: N.W. Kenya, Nandi and Teriki Hills, Elgon. 
crossleyi magnifica Rebel, 1914. Type-locality: Zaire, Nawambi-Irumu. 
= intermedia Joicey & Talbot, 1921. Type-locality: Zaire, Ituri 
Forest. 

Range: Zaire, E. and W. Uganda. 
crossleyi claudiae Rousseau-Decelle, 1934. Type-locality: Zaire, Katanga, 
Kafakumba. 

Range: Zaire in the Katanga district. 

Euxanthe (Euxanthe) wakefieldi (Ward) 

Euxanthe [Euxanthe) wakefieldi wakefieldi (Ward, 1873). Type-locality: E. Africa 
at Ribe, coastal hinterland, 
f. rubiginea Le Cerf, 1923. Type-locality: Tanzania, Nguru. 

Range: coastal forests of Kenya, Tanzania extending S. to Delagoa 
Bay, Malawi and inland to Zambia. 

Euxanthe [Euxanthe) madagascariensis (Lucas) 

Euxanthe [Euxanthe) madagascariensis (Lucas, 1842). Type-locality: Madagascar. 
Range: Malagasy Republic. 



94 V. G. L. van SOMEREN 

Euxanthe (Hypomelaena) trajanus (Ward) 

Euxanthe {Hypomelaena) trajanus trajanus (Ward, 1871). Type-locality: Cameroun. 
= schatzi Staudinger, 1885. Type-locality: Cameroun. 

Range: Cameroun and adjacent countries to W. of Congo River. 
trajanus vansomereni Poulton, 1929. Type-locality: Uganda, Mawakota 
District. 

Range: Uganda, central forests extending east to Mabira Forest. 
trajanus gabonicus Le Cerf, 1923. Type-locality: Gabon, Lambareni 

on Ogowe River. Status doubtful through lack of material, 
f. depuncta Le Cerf, 1923. Type-locality: Gabon, Ngemo on Ogowe 
River. 

Range: Gabon, Ogowe River. 
trajanus antonius Rousseau-Decelle, 1930. Type-locality: Zaire, Katanga, 
Kafakumba. 
Range: as type locality. 
trajanus nigeriae subsp. n. Type-locality: Nigeria, Ikom. 
Range: Nigeria. 

Euxanthe (Hypomelaena) tiberius Grose-Smith 

Euxanthe [Hypomelaena) tiberius tiberius Grose-Smith, 1889. Type-locality: Mom- 
basa area. 

f. tiberiella Strand, 1911. Type-locality: Tanzania, Amani. Range: 
coastal forests of Kenya and the Usambara Range, Tanzania. 

tiberius meruensis van Someren, 1935. Type-locality: Meru, Kenya. 
Range: East Mt Kenya, lower Meru forest and Meru Game Park. 



A BRIEF HISTORICAL REFERENCE 

Rothschild & Jordan (1898; 1900; 1903) published the results of their investigation 
into the genus Charaxes and its allies in their monumental work A Monograph 
of Charaxes and the Allied Prionopterous Genera. They employed the 'omnibus' 
genus Charaxes for all the African species with the exception of Palla and Euxanthe, 
which they dealt with in 1903. 

Aurivillius (1911), when dealing with the African Rhopalocera, arranged the 
species in groups under the genus Charaxes, retaining Palla and Euxanthe as distinct 
genera. 

Stichel (1939) gave a full list of the African 'Charaxidinae', supplying at the 
same time an exhaustive list of references, a monumental work in itself. He 
followed the general arrangement of Poulton (1926) but rearranged the groups 
and some of the species within them. 

Peters (1952) published his A Provisional Check-list of the Butterflies of the Ethiopian 
Region and, in the section on 'Charaxidinae', also followed Poulton (1926) and the 
general arrangement adopted by the British Museum (Natural History). Poulton 



REVISIONAL NOTES ON AFRICAN CHARAXES 95 

had divided the genus Charaxes into the Hadrodontiae and Leptodontiae based on 
the character of the fore wing costa. The Hadrodontiae have the costa coarsely 
serrate and include species No. 1-55 in my Synoptic List compared with the 
Leptodontiae with the costa finely serrate which contain the remaining species 
No. 56-119 of the Synoptic List. These 'subgenera' were divided into groups and 
subgroups but Peters placed Euxanthe before and Palla after Charaxes. 

In my Revisional Notes on African Charaxes, I have endeavoured to reassess 
the species and subspecies occurring in Africa and the adjacent islands, including 
Madagascar. I have not followed any accepted order or grouping but have left this 
to the Synoptic List, which broadly follows Peters (1952). Where I have advocated 
a departure from the hitherto accepted classification I have done so as a result of 
personal study of the imagines, as well as the early stages and foodplants, by 
rearing many broods from captive females of a great number of species. Thus I 
have endeavoured to make some sense of the 'black' Charaxes centred around 
Charaxes etheocles. 

Certain species appear to form compact groups, based on morphological characters; 
thus varanes, fulvescens and acuminatus all have characters in common and feed 
on Allophylus (Sapindaceae) and are now placed in the subgenus Stonehamia 
(Cowan, 1968 : 6), which replaces Hadrodontes Stoneham. 

On the evidence of the early stages and foodplants I have followed Rydon (1971) 
and raised both Palla and Euxanthe to subfamily status (Pallinae and Euxanthinae) , 
retaining the subgeneric name Hypomelaena Aurivillius for the two species with 
black undersides in the males, viz. trajanus and tiberius. 

When the early stages of the Charaxes doubledayi-mycerina group have been 
recorded the probability is that this group, too, will warrant subfamily status. 



Part I. 



p- 


197 line 13 




„ 38 




„ 39 


p- 


198 line 2 


p- 


205 „ 37 




.. 39 


p- 


206 line 4 


p- 


207 line 19 




„ 20 


p- 


220 line 32 




„ 36 




.. 4i 



CORRIGENDA TO PARTS I-IX 



for 'pythodorus' read pythodoris 

for 1925 read 1911-12 

after 'his' insert A Provisional 

after 'Check' insert hyphen 

after 'the' insert Butterflies of the 

after 'Ethiopian'' insert Region 

delete 'Butterflies'' 

for 'Cramer' read (Cramer) 

delete 'Charaxes'' insert Papilio Eques Achivus 

for '1776' read 1775-76 

for 'Check List' read check-list 

for 'Cramer' read (Cramer) 

ditto 

for 'PYTHODORUS' read PYTHODORIS 

for 'pythodorus' read pythodoris 

for 'pythodorus pythodorus" 1 read Pythodoris pythodoris 



9 6 V. G. L. van SOMEREN 

p. 221 Caption for 'PYTHODORUS PYTHODORUS' read PYTHODORIS 
PYTHODORIS 
Legend for 'pythodorus' read pythodoris 
p. 222 line 8 for 'pythodorus pythodorus' read pythodoris pythodoris 

,, io for 'pythodorus' read pythodoris 
p. 223 line 13 ditto 

,, 14 for 'pythodorus' read pythodoris 

„ 33 ditto 
p. 224 line 24 ditto 

,, 26 for 'pythodorus' read pythodoris 

,, 42 ditto 
p. 225 line 23 for 'pythodorus' read 'pythodoris 1 

„ 24 for 'pythodorus pythodorus' read pythodoris pythodoris 
p. 233 line 2 for sp. read ssp. 

25 after 'male' insert colon 
Index for 'pythodorus' read pythodoris 
PI. 12 Legend line 4 for 'Pythodorus pythodorus' and 'pythodorus' read pythodoris 

pythodoris and pythodoris 
PI. 13 Legend line 2 for 'pythodorus' read pythodoris 



Part II. 

p. 206 line 25 for '(Schultze)' read Schultze 

PI. 1 Legend line 2 for 'Cramer' read (Cramer) 
line 3 ditto 

„ 4 for 'Stoll' read (Stoll) 

„ 5 ditto 

„ 6 ditto 

PI. 2. Legend line 2 for 'Cramer' read (Cramer) 
PL 3 ditto 

PI. 4 ditto 

PI. 5 ditto 

PI. 6 ditto 

PL 7 ditto 



Part III. 

p. 47 after line 11 insert 3. The Charaxes etheocles (s.l.) Complex Part i 
line 12 delete '3' 
„ 15 delete 
„ 16 

„ 17 
p. 69 lines 32, 33 transfer to after line 18 
p. 70 after line 11 insert 3. The CHARAXES E THEOCLES (s.l.) Complex Part i 



REV1SIONAL NOTES ON AFRICAN CHARAXES 97 

p. 70 line 12 delete '3' 

,, 16 after 'Char axes' insert etheocles 

,, 17 after 'Jordan' insert as a subspecies of etheocles 
p. 74 line 27 delete 

lines 28-33 transfer to p. 97 below line 25 
p. 90 line 14 for '269' read 270 
p. 92 line 5 delete '1889' insert 1899 
p. 96 line 20 for 'Chriten' read Christon 
p. 97 after line 25 insert lines 28-33 from p. 74 
p. 98 line 30 delete 'Delagoa Bay, Zomba and Taveta' 

Part IV. 

p. 279 line 6 for 'Boisduval' read (Boisduval) 

,, 12 for 'Cramer' read (Cramer) 

„ 21 for 'Boisduval' read (Boisduval) 
p. 281 line 23 for 'Boisduval' read (Boisduval) 

„ delete Char axes insert Nymphalis 
p. 287 after line 10 insert Allotype female. Same data as holotype. 
p. 288 line 16 for '32' read 23 
p. 291 line 40 for ' citheronoid.es' read cithaeronoid.es 

p. 311 line 10 delete 'Kilimanjaro at Wasendo, 6000 ft. Type <J.' insert Type male. 
Type locality. Tanzania, Usambara Range, Magamba Forest nr 
Lushoto. 

,, 13 for 'Cramer' read (Cramer) 

,, 34 for 'Cramer' read (Cramer) 

,, 36 delete 'Charaxes' insert Papilio Eques Achivus 
p. 315 line 1 for 'CHARXAES' read CHARAXES 

,, 13 for 'Cramer' read (Cramer) 

,, 14 ditto 

p. 316 line 8 for 'Charaxes' read Charaxes 
PI. 9 legend line 2 for 'Cramer' read (Cramer) 
„ 5 ditto 

Part V. 

p. 77 line 16 After 'Complex' insert, Part 2 

p. 81 Map 1 Key, abbreviation for Charaxes is Ch., not 'C, also on Maps ^~7 and 

throughout text. 
p. 98 line 36 ditto 

p. 99 after line 29 insert 'Nymphalis ephyra Godart t.c. 
p. 101 line 33 for 'Godart' read (Godart) 
p. 114 line 10 for 'hollandi' read = hollandi 
,, 20 delete 'hollandi' insert ephyra 
p. 125 line 30 delete '25 : 42-43' insert 32 : 141-172 



9 8 V. G. L. van SOMEREN 

p. 132 Map 6 Key line 22 for 'Som' read Son 

,, 26 for 'figini' read fagini 
p. 133 after line 27 insert Charaxes chanleri Holland, 1896a : 262 [<$] 

line 28 for '1895 : 753' read 1896b : 753 [$] 
p. 165 Index for 'catachrous' read catochrous 
p. 166 ,, after line 16 insert fagini, 149 

Part VI. 

p. 199 line 8 for 'Dewitz' read (Dewitz) 

,, 12 for 'Cramer' read (Cramer) 

,, 18 for 'Drury' read (Drury) 

,, 28 for 'Dewitz' read (Dewitz) 

,, 28 abbreviations for Charaxes is Ch., not 'C throughout text and on 
Maps 
p. 212 line 36 for 'Cramer' read (Cramer) 
p. 214 line 4 for 'Cramer' read (Cramer) 

,, 27 for 'Cramer' read (Cramer) 

,, 29 for 'eques' read Eques 
p. 221 after line 23 insert Charaxes brutus andara Ward, Rothschild & Jordan, 

1900 : 435 
p. 222 line 40 for '(Cramer, 1779)' read (Cramer), 1779. 
p. 235 line 20 for ' 'quanzensis' read cuanzensis 
p. 236 line 19 for 'Drury' read (Drury) 
p. 238 line 32 for 'Drury' read (Drury) 
p. 239 line 20 ditto 

,, 22 delete 'Charaxes' insert Papilio Eques Achivus 
p. 247 line 22 for 'Drury' read (Drury) 

,, 23 ditto 

PI. 1 legend line 3 for 'Dewitz' read (Dewitz) 
PL 3 line 4 for 'Cramer' read (Cramer) 

PL 8 line 3 for 'Drury' read (Drury) 

PL 9 line 2 ditto 

PL 10 line 2 ditto 

Part VII. 

p. 183 line 7 for 'Cramer' read (Cramer) ; for 'hansalii' read hansali 
,, 9 ditto 

„ 25 ditto; for 'HANSALII' read HANSALI 

„ 26 for 'Cramer' read (Cramer) 
p. 184 line 5 ditto 

p. 190 line 14 for 'hansalii hansalii' read hansali hansali 
p. 191 caption line 1 for 'HANSALII' read 'HANSALI' 
lines 2-5 for 'hansalii' read hansali 
line 5 for 'hansalii' read hansali 



REVISIONAL NOTES ON AFRICAN CHARAXES 



99 



for 'hansalii' read hansali 
for 'hansalii' read hansali 
for 'Cramer' read (Cramer) 

ditto 
for 'antigua' read antiqua 
for 'hansaliV read hansali 
for 'hansalii hansalii'' read hansali hansali 
for 'hansalii' read hansali 
ditto 
ditto 
for 'Cramer' read (Cramer) 
for 'Cramer' read (Cramer) 
for e^wes read Eques 
for 'Hewitson' read (Hewitson) 
after 'adusta' insert aginga, 186 
for 'antiqua'' read antiqua 
for 'hansalii' read hansali 
PI. I Legend line 2 for 'Cramer' read (Cramer) 
PL 2 „ 2 ditto 

PI. 3 legend lines 9-1 1 for 'hansalii' read hansali 
PI. 4 legend line 2 for 'hansalii' read hansali 
2 for 'Cramer' read (Cramer) 

6 for 'Hewitson' read (Hewitson) 

7 ditto 

6 for 'Drury' read (Drury) 

7 ditto 



p. 


192 line 35 


p- 


193 line 5 




„ 19 




„ 20 




„ 26 


p- 


194 line 2 




„ 3 




» 5 




„ 12 




» 14 




„ 16 




„ 18 


p. 


195 line 4 


p- 


207 line 41 


p. 


225 Index 


p- 


226 index 



PL 5 
PI. 8 

PI. 9 



Part VIII 




p. 217 line 


9 


>> 


11 


" 


20 


»> 


22 


>> 

„ 

p. 218 line 


27 


3i 

2 


p. 219 line 

>> 


15 
16 


21 


>9 


23 


\ 220 captior 



for 'Dewitz' read (Dewitz) 

for 'thysii' read thysi; for 'C read Ch. 

for 'Cramer' read (Cramer) 

for 'Hewitson' read (Hewitson) 

for 'C read Ch. 

ditto 
for 'Schultz' read Schultze 
for 'mccleeryi' read maccleeryi 
for 'Dewitz' read (Dewitz) 
for 'Dewitz' read (Dewitz) 

ditto 
for 'THY SW read THYSI, for 'C read CH. 
for 'thysii' read thysi 
line 4 for 'thysii 1 read £Aysi 



ioo V. G. L. van SOMEREN 

p. 222 line 12 for '1925' read 1926 

„ 31 for 'thysii' read thy si 

„ 32 for 'thysii' read thysi 

,, 38 for '1925' read 1926 
p. 227 line 33 for '211' read 271 
p. 228 line 14 after Cottrell). delete 

line 13 for 'Lisombe' read Lisombo 

,, 15 delete 

,, 16 delete 'Malawi and adjacent' 
p. 235 line 29 for 'Cramer' read (Cramer) 
p. 240 line 3 for 'Cramer' read (Cramer) 

,, 4 for 'Cramer' read (Cramer) 

„ 30 for 'C read CH. 

for 'Hewitson' read (Hewitson) 
p. 241 line 23 for '1865' read 1859 
p. 245 line 14 for 'Ungemache' read Ungemach 

„ 20 for *C\ read Ch. 

„ for 'Hewitson' read (Hewitson) 

,, 21 for '1865' read 1859 
p. 253 line 24 for 'barnesi' read barnsi 

,, 26 for 'barnesi'' read barnsi 
p. 254 line 38 for 'barnesi' read barnsi 

,, 39 for 'barnesi' read barnsi 
p. 255 line 3 for 'C read CH. 

for 'MCCLEERYI' read MACCLEERY1 
p. 257 line 21 for 'mccleeryi' read maccleeryi 

,, 27 for 'mccleeryi' read maccleeryi 
p. 258 line 31 for 'Kleilland' read Keilland 

,, 42 for 'mccleeryi' read maccleeryi 

„ 43 for 'mccleeryi' read maccleeryi 
p. 260 line 32 delete 'ssp. n.' insert van Someren 

after line 33 insert See Charaxes xiphares kilimensis van Someren, 1969 : 82 
lines 34-46 delete 
p. 261 delete 
p. 262 lines 2-6 delete 

after line 9 insert Charaxes imperialis ludovici Rousseau-Decelle, Stichel, 

1939 = 452 
line 12 for 'race,' read race, and delete remainder of sentence. 
„ 14 delete '(Grahame coll.)' (MNHN, Paris) 
„ 23 delete 'ssp.n.' insert van Someren, 1969. 
p. 264 Index for 'barnesi' read barnsi 

for 'mccleeryi' read maccleeryi 
for 'lecerfi, 232' read lecerfi, 222 
for 'thysii' read thysi 
PI. 1 Legend line 4 for 'hiblderandti Dewitz,' read hilderbrandti (Dewitz), 



REVISIONAL NOTES ON AFRICAN CHARAXES 

PL i Legend line 5 for 'Dewitz' read (Dewitz) 

„ 8 for 'thysW read thysi 

„ 11 for 'lecer\' read lecerfi 

„ 13 ditto 

PI. 7 Legend line 7 for 'Cramer' read (Cramer) 

,, 9 ditto 

PL 8 Legend line 8 for 'Hewitson' read (Hewitson) 

,, 9 ditto 

„ 10 ditto 

,, 11 ditto 

PL 11 Legend line 8 for 'barnesi' read barnsi 

PL 12 legend line 7 for 'mccleeryi' read maccleeryi 

,, 9 ditto 

,, 14 delete '(I. Grahame)' insert (MNHN, Paris) 

,, 15 delete 



Part IX. 




P- 


417 


line 8 






>> 


10 






it 


16 






1 y 


22 
25 






>> 


28 






tt 


31 






a 


35 
39 


P- 


418 


line 


2 






) > 


3 
16 

18 


P- 


424 


line 


35 






>> 


36 


P- 


425 


line 


13 


P- 


444 


line 


25 


P. 


452 


line 26 






II 


28 


P- 


460 


line 


13 


P- 


462 


line 


3 






t$ 


23 






ji 


29 






>> 


30 


P- 


403 


line 


19 


P- 


466 


line 


19 



for 'Trimen' read (Trimen) 

for 'Godart' read (Godart) 

for 'Drury' read (Drury) 

for 'Westwood' read (Westwood) 

for 'Godart' read (Godart) 

for 'Drury' read (Drury) 

for 'Stoll' read (Stoll) 

for 'Godart' read (Godart) 

for 'Cramer' read (Cramer) 

for 'Trimen' read (Trimen) 

for 'Trimen' read (Trimen) 

ditto 
delete 'Char axes' insert Nymphalis 
for 'Trimen' read (Trimen) 

ditto 
for 'Godart' read (Godart) 
for 'Westwood' read (Westwood) 
for 'Godart' read (Godart) 
delete 'Charaxes 1 insert Nymphalis 
insert Nymphalis before nesiope 
for 'his' read Rochat 
for 'Plantrou' read Rochat 
for 'Godart' read (Godart) 

ditto 
for 'Drury' read (Drury) 

insert Nymphalis after 'Fapilio'; for 'phaleratus' read Phalerakis; 
for '1872' read 1782 



V. G. L. van SOMEREN 



p- 


467 line 33 


for 'Stoll' read (Stoll) 


p- 


470 line 3 


insert PL 15, fig. 168 


p- 


472 line 3 


delete 168 insert 167 


p- 


476 line 8 


for 'Cramer' read (Cramer) 


p- 


480 line 24 


for '(Cramer, 1764).' read (Cramer), 1764. 




„ 28 


for '(Mabille, 1876).' read (Mabille), 1876. 


p- 


483 line 44 


for 'lactitinctus Karsch' read lactetinctus ungemachi Le Cerf 




„ 46 


for 'lactitinctus' read lactetinctus 


p- 


486 Index 


for lactitinctus read lactetinctus 


p- 


487 Index 


for 'ungemachi' read ungemachi 


PI 


. 1 Legend line 2 for 'Trimen' read (Trimen) 






,,4 for 'Trimen' read (Trimen) 






,, 6 ditto 






,, 7 ditto 






„ 8 ditto 


PI 


. 9 Legend line 4 for 'Godart' read (Godart) 






„ 5 ditto 






,, 6 ditto 


PI 


. 11 Legend 


line 14 for 'Drury' read (Drury) 
„ 15 ditto 
,, 16 ditto 


PI 


. 12 Legend 


line 4 ditto 


PI 


. 13 Legend 


line 8 for 'Stoll' read (Stoll) 
,, 9 ditto 

,, 10 for 'Godart' read (Godart) 
„ 11 ditto 


PI 


. 14 Legend 


line 4 ditto 
„ 5 ditto 
,,6 for 'thomassius' read thomasius 


P] 


. 15 Legend 


line 9 delete '& 168' and one $ symbol 

ifter line 9 insert Fig. 168 candiope thomasius Staudinger & Schatz $ 

(Sao Thome) 


PI 


. 17 Legend 


line 2 for 'Cramer' read (Cramer) 


PI 


.18 Legend 


line 14 for 'lactitinctus' read lactetinctus 
,, 16 ditto 



ADDENDA 

Since the submission of the typescript, photographs and maps for Part X of 
my Revisional Notes on African Charaxes, several new species and subspecies have 
been added to the taxa dealt with in this series. The following addenda include 
the data, both published and unpublished, which have come to my notice up to 
June 1974. The opportunity is also taken to elevate certain taxa within the 
'black Charaxes' complex to specific status and to re-name certain taxa which have 
been found to be homonymous. I am indebted to Mr T. G. Howarth, Dr A. H. B. 
Rydon and to Mr C. F. Huggins for much help with the preparation of these addenda 



REVISIONAL NOTES ON AFRICAN CHARAXES 103 

and also wish to thank Messrs W. H. Henning, S. C. Collins and R. S. White for 
the loan of material and coloured photographs. 

FURTHER DESCRIPTIONS AND NOTES ON SPECIES AND SUBSPECIES 

OF CHARAXES 

Charaxes fulvescens (Aurivillius) 

Charaxes fulvescens (Aurivillius); van Someren, 1963 : 210. 

Dr M. Condamin of I.F.A.N., Dakar, has recently drawn my attention to the fact 
that Charaxes fulvescens extends into Senegal, but it is here represented by a distinct 
subspecies. Through the kindness of Dr Condamin I have been able to examine 
a series of three males and two females. 

Charaxes fulvescens senegala subsp. n. 

Male. Fore wing length 43-45 mm. Apex bluntly pointed, outer margin slightly concave 
in spaces 3-5. Hind wing margin rather rounded with a short blunt tail, 5 mm in length 
at vein 4. Upperside. Ground colour darker than in nominate race. Fore wing, basal area 
more fulvous but paler at the base of the hind wing, but both with a slight greenish tinge. 
Discal zone more rufescent, especially in hind wing where it borders on the darker distal portion 
of the wings. Fore wing with a series of large rufous-tawny spots in the post discal line, extend- 
ing from the subcosta to xb flanked by a more obscure series in the submarginal zone. Two 
large contiguous spots present beyond the cell ; flanked by dark spots at apex of cell and the 
subbasal areas of spaces 4-2. Hind wing, basal area as fore wing with a strong rufescent zone 
bordering the proximal edge of the dark border carrying a series of somewhat rounded obscure 
dark spots. Edge of wing slightly rufous. Underside. Fore wing, basal area dark greyish 
brown with a few fine black lines and dots towards the base, accentuated in the discal line by a 
blacker line edged distally with whitish. This line is angled at vein 6, then curves to the hind 
margin and continues in the hind wing where it is almost straight from mid-costa to the hind 
angle. The distal portions of both fore and hind wing have a slight satiny sheen. A large 
dark ocellus in subcosta of space 6 followed by very obscure spots in the postdiscal area. 

Female. Fore wing length 50-51 mm. Similar in markings to male. 

Holotype <§, Senegal: For£t classee Santiaba-Mandjak, 4.XL1965 (M. Condamin) 
(MNHN, Paris). 

Allotype $, Senegal: Foret classee Santiaba-Mandjak, 19.X.1962, sur banane 
fermentee (Mission IFAN en Basse Casamance) (MNHN, Paris). 

Paratypes. 1 <$, same data as holotype but 13.X.1962 (IFAN, Senegal); 1 $, same 
data as allotype but 18.X.1962 (BMNH); 1 <J, Foret classee de Tobor, 21.xi.1961, 
sur banane fermentee (Mission en Basse Casamance) (BMNH) . 

Charaxes octavus Minig 
(PI. 18, figs 137, 138) 

Charaxes octavus Minig, 197 1 : 269. 
Charaxes patergodarti Neidhoefer, 1972 : 5. 

The description of this new species is based on a single male specimen taken at 
Bangui, Central African Republic. According to Neidhoefer his specimen, which is 



104 V. G. L. van SOMEREN 

also a male from Bangui, and which is possibly the same specimen as that described 
by Minig, is a large insect similar in coloration to Ch. eudoxus and Ch. lucretius on 
the upperside. Its underside is a unique combination of both species but is closer 
to lucretius. 

Charaxes ansorgei rydoni van Someren 
(PI. 18, figs 139, 140) 
Charaxes ansorgei rydoni van Someren, 1967 : 309. 

The description of this distinctive race was based on males only, but the female 
has since been taken by Mr Ivan Bampton in the Magamba Forest, Lushoto, 
Tanzania. I am indebted to Mr W. Henning for colour photographs of this sex. 

Female. Upperside. Fore wing length 47 mm. Basal area chestnut with obscure black 
marks in the mid cell and inward to its end where they merge into the black distal half of the 
wing, which is crossed by a series of spots, large and whitish in area 1 and 2, more ochreous in 
3-4 in subcostal region. Postdiscal spots discreet and more strongly orange-ochreous, but 
conjoined with the discal spots in 3—4. Outer half of wing black but with conspicuous orange 
ochreous spots on the margin. Hindwing, basal area chestnut, shading to greyish on the 
inner fold. Discal band white, mainly broad but abruptly narrowing towards costa on proximal 
edge, but here indicated by obscure whitish marks. Outer border black, narrowing at anal 
angle. Admarginal orange triangular marks conspicuous, becoming linear at anal angle. 
Anal angle with two blue spots, lower one large. Margin serrate with long tapering pointed 
tails at 3 and 6. Underside. More boldly marked than in other races; the general pattern 
somewhat similar to that of the male but markings bolder and more conspicuous, the purple 
anal spot and one above very bold. 

Neallotype $, Tanzania: Usambara Mts, Lushoto, Magamba, 6000 ft, 4.iii.iQ73 
(/. Bampton) (W. H. Henning Coll.). 



Charaxes eudoxus lucyae subsp. n. 

(PI. 18, figs. 141-144) 

Female. Length of fore wing 44 mm and 30 mm from mid-costa to hind angle. Upperside* 
Fore wing, basal area bright chestnut; end of cell with slightly curved black lines; discal spots 
black, large at bases of spaces 4 and 3, slightly indicated in upper part of 2; beyond, towards 
costa two black spots, irregular in shape. Discal band orange, widest at the hind margin and 
tapering to subapex; the three subapical spots rounded and distinct. Border of wing black, 
extending along veins; interspaces orange, extending to apex. Hindwing, basal area orange- 
chestnut, shading to greyish, tinged at base of inner fold, both shading distally into the orange- 
yellow band which is darker orange-yellow to above the anal angle. This is followed by a 
broad black band, widest at upper angle and extending to anal angle which has two blue spots. 
Border orange and broad margin serrate with black edge; long thin pointed tails at veins 4 
and 6. Underside. Very orange. Fore wing, ground colour bright chestnut from base to 
subapex, strongly marked with black oblong spots, circular at base, more irregular, elongate 
in cell and linear at subapex, all strongly outlined in white. Post discal bar wide at base and 
triangular, becoming a series of double lunate marks. The bar is white inwardly, shaded 
with orange distally to margin and with conspicuous black marks in hind angle and with 
violet spots in 4-5. Margin of wing orange, the tips of the veins indicated as upperside. Hind 



REVISIONAL NOTES ON AFRICAN CHARAXES 105 

wing, bright chestnut at base widening in post discal zone; the base with black lines forming 
elongate triangles on inner fold and lower disc, all outlined in white. Discal bars white, double 
at costa, then going straight to inner fold; the double lines with paler orange-chestnut between. 
Post discal zone bright chestnut, widest towards upper angle, then tapering towards anal angle 
which carries double rounded conjoined spots with bluish purple centres. The band with 
indistinct shading centrally, but margined distally with black, accentuated distally with white. 
Edge of wing orange, accentuated at margin with black. 

Holotype 2, Tanzania: Usambara Mts, Magamba Forest, Lushoto, 6000 ft, 
4.111.1973 (/. Bampton). To be deposited in BMNH. 

Paratype2, Tanzania: Usambara Mts, Magamba Forest, Lushoto, 5000 ft, ii. 1974 
(S. C. Collins). Deposited in BMNH. 

Charaxes cithaeron cithaeron Felder 
2 aberration whitei ab. n. 

(PL 19, figs 145, 146) 

Differs from the nominate female by having the hind wing upperside discal area pale ochreous 
instead of bluish white. The ground colour of both wings on upper- and underside paler 
than normal but this may be due to the worn and tattered condition of this specimen. I 
have been unable to trace any record of a similar specimen in collections or in the literature. 

Holotype 2, South Africa: Natal, Eshowe, 8.V.1971 (R. S. White) B.M.1974-332. 

Charaxes pythodoris davidi Plantrou 

Charaxes pythodorus davidi Plantrou, 1973 : 269. 

Apparently this newly described subspecies from Ivory Coast is smaller and 
differs in shape and markings from both the nominate subspecies and subsp. occidens 
van Someren described from Central African Republic (French Congo) and Nigeria. 

Charaxes hildebrandti gillesi Plantrou 

Charaxes hildebrandti gillesi Plantrou, 1973 : 274. 

Apparently this newly described subspecies occurs in the Ivory Coast and Ghana 
and in both sexes may be distinguished from the nominate subspecies by the wider 
creamy white median band on the upper and underside of both fore and hind wings. 

Charaxes usambarae van Someren & Jackson 
9 form collinsi forma n. 

(PL 19, figs 147, 148) 

Upperside. Differs from the nominate female in that the fore wing discal bar is wider 
at the base, forming a triangle from the hind margin to the cell ; the spots above angles on the 
distal side, the uppermost slightly inset. The subapical spots white; the upper three in line, 
the two below smaller and inset. Two indistinct whitish spots at hind angle. Hindwing, 



io6 V. G. L. van SOMEREN 

discal bar wider and more diffuse on margins. Submarginal linear spots more distinct, so also 
the marginal lunules. Edge of wing slightly more serrate; tails longer and thinner. Underside. 
Fore wing, ground colour paler, darker on border. Base with three rounded black spots outlined 
in white followed by a wavy black line in mid cell, and a similar mark beyond, black lines 
at bases of spaces i and 2 similarly outlined. Discal white bar narrower than upperside and 
less distinct, the spot in space 1 only joined slightly to post discal spot which is represented 
above by indistinct dyslegnic spots, the upper ones representing the white subapical spots of 
upper side; the whole row distally bordered by dark obscure marks, most distinct at hind angle. 
Hind wing much as in nominate form but with median band narrower, distally outlined by 
darker zigzag line. Submarginal line as in upperside but less distinct, ending in three spots in 
anal angle. Marginal border less distinct than upperside. 

Holotype $, Tanzania: Usambara Mts, Amani. Bred on Albizzia sp. by African 
collector for S. Collins, x. 1973. Deposited in BMNH. 

Charaxes martini van Someren 
(PI. 19, figs 149, 150) 

Charaxes martini van Someren, 1966 : 96 [<J]. 
Charaxes martini van Someren, 1974 : 483 [§]. 

This hitherto elusive species appears to be confined to the higher ground of 
Mlange Mt, especially near the Malosa Stream, in Malawi. When first described it 
was known only from the male and later a very damaged female was described. A 
female specimen in good condition has since been taken in the type-locality by Dr 
C. H. McCleery who has kindly supplied the photographs which are reproduced here. 
The male is characterised by the diffuse greenish subapical spots of the fore wing 
and the conspicuous greenish band in the discal portion of the hind wing, followed 
by a complete row of white spots; the margin strongly marked with reddish and 
greenish markings. The underside is strongly marked on a silvery ground by 
black lines. The female is also boldly marked both on the upper- and undersides. 
The pattern is very similar to that of the male. The upperside median bar is 
slightly tinged with yellowish and is slightly more tinged with greenish on the lateral 
edging of the hind wing. The hind wing underside is rather silvery with a greenish 
post discal bar. The tails are longer than those of the male. 

Charaxes mafuga van Someren 

(PI. 19, figs 151, 152) 

Charaxes mafuga van Someren, 1969 : 97. 

This species was originally described from male specimens taken in the Mafuga 
area of Kigezi in S.W. Uganda. The female had not been authentically identified 
until it was bred from a larva found by Mr I. Bampton in the same area. 

Female. Larger than the male but with the same distinctive underside. Fore wing 
length 37-5 mm. Upperside. Fore and hind wing ground colour black, slightly more brownish 
towards base of fore wing. Fore wing, discal line of spots from beyond the cell blue, almost 
straight except for spot in apex of cell, gradually increasing in size to hind margin; post discal 
spots of subapex in a line, followed by spots of increasing size, those towards the hind margin 



REVISIONAL NOTES ON AFRICAN CHARAXES 107 

larger and contiguous with the discal spots. Hind wing, blue band almost straight on inner 
edge, single on the costa but fused beyond but not reaching the inner fold. Submarginal 
marks linear and blue and distinct, double at anal angle. Admarginal marks linear to upper 
tail; edge of wing bluntly serrate, bluish olive in colour. Tails; upper slightly spatulate, lower 
shorter and pointed. Underside. Fore wing, greyish brown in basal area, more brownish 
distally. Base crossed by fine black lines ; disc crossed by greyish white bar, widest towards the 
end of cell, accentuated distally by darker brown separating it from the whitish post discal 
series of silvery grey dyslegnic spots which are conjoined with those of the subapex. This 
bar is accentuated distally by a series of brownish grey marks widest towards the hind angle, 
this in turn is accentuated by silvery grey, which is more brownish towards the outer margin. 
Hind wing, basal area silvery grey, paler towards the inner fold, crossed in the subbase by a 
whitish bar. Discal bar distinct, narrower and zigzag in mid area, widening to the costa but 
here accentuated by the darker brown post discal band which carries a series of crescentic 
marks distally. Border pale grey with a series of white linear marks outwardly edged with 
blue; margin narrowly edged with orange that has a greenish tinge. 

Neallotype $, Uganda, Ruhiza, Impenetrable Forest, 8000 ft, 28.ix.1972 
(J. Bamptori) B.M. 1972-571. 

A FURTHER NOTE ON THE 'BLACK' CHARAXES COMPLEX 

As a result of breeding from known females by Mr I. Bampton and Mr W. H. 
Henning in some cases, and on the evidence revealed by the examination of the 
male genitalia by Mr G. A. Henning and Dr A. H. B. Rydon in most cases, the 
raising in status of the names listed below is now suggested. 

It is clear from their original descriptions that pseudophaeus and chintechi were 
proposed as infrasubspecific names, and have thus been strictly unavailable under 
the International Code of Zoological Nomenclature until now; protomanica, proposed 
for a form after i960, and vansoni, proposed for a form of a subspecies, have also 
been hitherto unavailable under the Code. These four names thus become available 
now for the first time and are therefore marked 'sp. n.' in the list below. References 
are given to their original descriptions as forms, and their type-material is that 
cited at the time of these original descriptions. The remaining six names have had 
previous availability as species-group names. 

Charaxes nyikensis van Someren stat. n. 
Char axes alpinus nyikensis van Someren, 1966 : 85. 

Charaxes pseudophaeus van Someren sp. n. 

Charaxes manica 9 f- pseudophaeus van Someren & Jackson, 1957 : 46. 

Charaxes manica $ f. pseudophaeus van Someren & Jackson; van Someren, 1966 : 89. 

Charaxes chintechi van Someren sp. n. 

Charaxes manica $ f. chintechi van Someren & Jackson, 1952 : 270. 

Charaxes manica ^ f. chintechi van Someren & Jackson; van Someren, 1966 : 90. 

Charaxes protomanica van Someren sp. n. 
Charaxes manica § f. protomanica van Someren, 1966 : 91. 



io8 V. G. L. van SOMEREN 

Charaxes pondoensis van Someren stat. n. 
Char axes ethalion pondoensis van Someren, 1967 : 285. 

Charaxes phaeus Hewitson stat. rev. 

Charaxes phaeus Hewitson, 1877 : 82. 

Charaxes viola phaeus Hewitson; van Someren, 1969 : 136. 

Charaxes vansoni van Someren sp. n. 

Charaxes viola phaeus $ f. vansoni van Someren & Jackson, 1957 : 43. 

Charaxes viola phaeus $ f- vansoni van Someren & Jackson; van Someren, 1969 : 137. 

Charaxes variata van Someren stat. n. 
Charaxes viola variata van Someren, 1969 : 144. 

Charaxes loandae van Someren stat. n. 
Charaxes viola loandae van Someren, 1969 : 144. 

Charaxes brainei van Son stat. n. 

Charaxes viola brainei van Son, 1966 : 3. 

Charaxes viola brainei van Son ; van Someren, 1969 : 147. 

SOME REPLACEMENT NAMES AND ADDITIONAL INFORMATION 

ON CERTAIN TAXA 

Charaxes hansali kulalae nom. n. 

Charaxes hansalii kulalensis van Someren, 1971 : 192. 

The name kulalae is proposed to replace the name kulalensis van Someren in the 
combination Charaxes hansalii kulalensis van Someren, 1971 (Part VII), which is a 
junior primary homonym of kulalensis van Someren in the combination Charaxes 
acuminatus kulalensis van Someren, 1963 (Part I : 217). 

Charaxes jahlusa kigomaensis nom. n. 

Charaxes jahlusa kigoma van Someren, 1974 : 423. 

The name kigomaensis is proposed to replace the name kigoma van Someren in 
the combination Charaxes jahlusa kigoma van Someren, 1974 (Part IX), which is a 
junior primary homonym of kigoma van Someren in the combination Charaxes 
smaragdalis kigoma van Someren, 1964 (Part II : 219). 

Charaxes anticlea suna nom. n. 

Charaxes anticlea reducta van Someren, 1971 : 214. 

The name suna is proposed to replace the name reducta van Someren in the 
combination Charaxes anticlea reducta van Someren, 1971 (Part VII) which is a 



REVISIONAL NOTES ON AFRICAN CHARAXES 109 

junior primary homonym of reducta Rothschild in the combination Charaxes xiphares 
reducta Rothschild, 1929 (Part II : 189). 

Charaxes ethalion nyasicus nom. n. 

Charaxes ethalion nyasana van Someren 1967 : 286. 

The name nyasicus is proposed to replace the name nyasana van Someren in the 
combination Charaxes ethalion nyasana van Someren, 1967 (Part IV) which is a 
junior primary homonym of nyasana Butler in the combination Charaxes nyasana 
Butler, 1895 (Part VII : 206). 

Comparatively recent examination (May, 1967) of additional material indicates 
that this subspecies extends into S. W. Katanga (see Part IV : 300, Group 3, 
Region 2). 

Charaxes dilutus miotoni nom. n. 

Charaxes dilutus ngonga van Someren, 1974 : 442. 

The name miotoni nom. n. is proposed to replace the name ngonga van Someren 
in the combination Charaxes dilutus ngonga van Someren, 1974 (Part IX), which is 
a potential junior primary homonym of ngonga van Someren in the combination 
Charaxes berkeleyi van Someren & Jackson $ form ngonga van Someren 1969 
(Part V : 80). 

Charaxes thy si Capronnier 

Charaxes thysii Capronnier; van Someren, 1972 : 221. 

Apparently, according to Dr A. H. B. Rydon (personal communication) , there is a 
female specimen of this species in the Musee Royal de l'Afrique Centrale, Tervuren, 
Belgium. 

Charaxes ludovici Rousseau-Decelle 

Charaxes ludovici Rousseau-Decelle, 1933 : 271. 

Mr Howarth has drawn my attention to the confusion regarding this name, which 
was based on a specimen from Lake Nyassa. Stichel (1939 : 452) placed this 
taxon as a subspecies of Charaxes imperialis Butler, and in Part VIII (p. 227) I 
wrongly assigned to this combination specimens taken by C. B. Cottrell at 
Mwinilunga, Zambia. Later in the same part (p. 262) I correctly placed the true 
ludovici from Lake Nyassa as a subspecies of Charaxes xiphares. The specimens 
taken by Cottrell in Zambia thus require a new name, as follows. 

Charaxes imperialis lisomboensis subsp. n. 

[Charaxes imperialis ludovici Rousseau-Decelle; van Someren, 1972 : 227. Misidentincation.] 

The name lisomboensis is proposed for the subspecies I misidentified in Part VIII 
as Charaxes imperialis ludovici Rousseau-Decelle. 



no V. G. L. van SOMEREN 

The male and female specimens described and figured in 1972 are here designated 
the holotype and allotype respectively. It should be noted that the remarks made 
on p. 229 lines 14 and 15 about these specimens being atypical should be deleted and 
also the reference to Malawi on line 16 (see Corrigenda). 

Holotype <$, Zambia: Mwinilunga, Lisombo River (C. B. Cottrell). 

Allotype $, same data as holotype. Types in C. B. Cottrell collection to be 
deposited in BMNH. 

Range. Zambia. 

Charaxes superbus Schultze 

Charaxes superbus Schultze; Plantrou, 1965 : 30 [$]. 
Charaxes superbus Schultze; van Someren 1974 : 436. 

When dealing with this species the reference to the description of the female 
by Monsieur J. Plantrou was unfortunately omitted. 



Charaxes lydiae Holland 

Charaxes lydiae Holland, 191 7 : 18. 
Charaxes lydiae Holland; Darge, 1973a : 51. 
Charaxes lydiae Holland; van Someren, 1974 : 4& 1 - 

Since writing Part 9 of these Revisional Notes Monsieur P. Darge has given an 
account of the discovery of this hitherto very rare species at Yaounde, Cameroun 
in sufficient numbers to enable him to describe the variation within the species. 



Charaxes eudoxus musakensis Darge 

Charaxes eudoxus musakensis Darge, 1973& : 2 9- 

This interesting subspecies has been described from Mt Cameroun in the Cameroun. 

Charaxes richelmanni Rober 

Charaxes richelmanni Rober; van Someren, 1970 : 236. 
Charaxes richelmanni Rober; Darge, 19736 : 26 [$]. 

Since this species was dealt with by me in 1970 the female has been described 
by Monsieur P. Darge from Mt Kala, Cameroun. 



SYNOPTIC LIST OF AFRICAN CHARAXES, 
EUXANTHE AND PALL A 

As the final sequence given below differs from the sequence in earlier parts (see 
p. 94), references are given in square brackets to the part number (Roman) and 
page numbers where each species is dealt with in the text. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



in 



CHARAXINAE Doherty 

CHARAXES Ochsenheimer 
Subgenus STONEHAMIA Cowan 11 



varanes-group 

i. varanes (Cramer) [IX : 476] 
v. varanes (Cramer) 

austrinus Rothschild 
v. vologeses (Mabille) 

brachycauda Le Cerf 
v. bertrami Riley 
v. defulvata Joicey & Talbot 

2. fulvescens (Aurivilhus) [I : 210] 

f. fulvescens (Aurivilhus) 

/. monitor Rothschild 

/. saperanus Poulton [not examined] 

/. senegala van Someren 

3. acuminatus Thurau [I : 211, V : 77] 

a. acuminatus Thurau 



a. vumba van Someren 
a. mlanje van Someren 
a. cottrelli van Someren 
a. nyika van Someren 
a. usambarensis van Someren 
a. shimbanus van Someren 
a. teitensis van Someren 
a. oreas Talbot 
a. kulalensis van Someren 
a. stonehami Jeffery 
a. kigezia van Someren 
a. obudoensis van Someren 
balfouri Butler [Not dealt with] 
analava Ward [IX : 472] 



Subgenus CHARAXES Ochsenheimer 



candiope-gvoup 

6. candiope (Godart) [IX : 467] 

c. candiope (Godart) 

viridicostatus Aurivilhus 
c. velox Grant 
c. thomasius Staudinger & Schatz 

7. antamboulou Lucas [IX : 472] 

8. cowani Butler [IX : 471] 
cynthia-group 

9. protoclea Feisthamel [VII : 202] 
jp. protoclea Feisthamel 

aeson Herrich-Schaffer 
p. protonothodes van Someren 
var. ablutus Schultze 
var. maculata Strand 
var. marginepunctata Holland 
var. nigropunctata Neustetter 
var. sinuosa Rousseau-Decelle 
p. nothodes Jordan 
p. catenaria Rousseau-Decelle 
var. bifida Rousseau-Decelle 
var. kafakumbana Rousseau- 
Decelle 
var. parcepicta Rousseau-Decelle 
var. mutschatschana Rousseau- 
Decelle 



12. 



p. azota (Hewitson) 

calliclea Smith 

nyasana Butler 

f. aequidistans Gaede 
boueti Feisthamel [VI : 224] 
b. boueti Feisthamel 
b. ghanaensis Rousseau-Decelle & 

Johnson 
b. macclouni Butler 

flavescens Lanz 
b. rectans Rothschild & Jordan 

b. alticola Grunberg 
lasti Grose-Smith [VI : 232] 

centralis Neustetter 
cynthia Butler [V : 150] 

c. cynthia Butler 

lysianassa Westwood 

guineensis Le Moult 
c. cameroonensis van Someren 

f . albofascia Le Cerf 

ab. cizeyi Lathy 

f. angusticlavius Rousseau-Decelle 
c. kinduana Le Cerf 

mawamba Grunberg 
c. propinqua van Someren 
c. parvicaudatus Lathy 



• The compact group varanes-fulvescens-acuminatus, usually placed in the genus Charaxes, was 
separated off by Stoneham under the name Hadrodontes but, as Cowan (1968) points out, the name is 
unavailable because no type-species was designated as required by Article 13 (b) of the Code. Cowan 
proposed Stonehamia as an alternative name, citing Papilio varanes Cramer as the type-species; the 
name can be used as a subgenus. 



V. G. L. van SOMEREN 



c. sabulosus Talbot 

f. aurantiaca Rousseau-Decelle 
c. mukuyu van Someren 
fucref »'as-group 

13. lucretius (Cramer) [VII : 194] 

I. lucretius (Cramer) 

lucida Le Cerf 
/. inter medius van Someren 

albofascia Le Cerf 

caliginosa Le Cerf 
I. maximus van Someren 

babingtoni Stoneham 
/. lemosi Joicey & Talbot 

14. octavus Minig [X : 103] 

patergodarti Neidhoefer 

15. odysseus Staudinger [VII : 199] 

16. lactetinctus Karsch [VIII : 232, IX : 

483] 
/. lactetinctus Karsch 
I. busogus van Someren 

f . jacksonianus van Someren 
/. ungemachi Le Cerf 
f. brunneus Carpenter 
jasius-gioup 

17. jasius (Linnaeus) [I : 201] 

j. jasius (Linnaeus) 
j. epijasius Reiche 

var. maculatus Suffert 

ab. murina Le Cerf 

ab. feisthameli Le Cerf 

var. melas van Someren 

f. liberiae Le Cerf 
j. harrisoni Sharpe 

f. harrisoni Sharpe 

f. saturnalis van Someren 
j. pagenstecheri Poulton 
j. saturnus Butler 

var. laticinctus Butler 
j. brunnescens Poulton 

18. pelias (Cramer) [I : 205] 

19. hansali Felder [VII : 190] 

h. hansali Felder 

h. baringana Rothschild 

h. kulalae van Someren 

kulalensis van Someren 
h. arabica Riley 

20. castor (Cramer) [VII : 184] 

c. castor (Cramer) 

ab. aginga Stoneham 

ab. antiqua Le Cerf 

ab. flavimarginalis Stoneham 

c. flavifasciatus Butler 
orientalis Lanz 
var. reimeri Rothschild 



c. arthuri van Someren 

c. comoranus Rothschild 

21. brutus (Cramer) [VI : 214] 

b. brutus (Cramer) 

b. angustus Rothschild 

b. Junius Oberthur 

b. somalicus Rothschild 

b. alcyone Stoneham 

b. natalensis Staudinger & Schatz 

b. antiquus Joicey & Talbot 

22. andara Ward [VI : 221] 

23. ansorgei Rothschild [IV : 303] 

a. ansorgei Rothschild 

a. jacksoni Poulton 

a. ruandana Talbot 

a. kungwensis van Someren 

a. levicki Poulton 

a. kilimanjarica van Someren 

a. rydoni van Someren 

24. phoebus Butler [VI : 212] 

25. pollux (Cramer) [IV : 311] 

p. pollux (Cramer) 

ab. subalbescens Hall 
var. ongeus Stoneham 
var. bungense Stoneham 

jp. geminus Rothschild 
zingense Stoneham 

p. maua van Someren 

p. gazanus van Someren 

26. druceanus Butler [I : 228] 

d. druceanus Butler 
d. tectonis Rothschild 
d. obscura Rebel 

kivuanus Jordan 

cryanae Le Cerf 
d. septentrionalis Lathy 

var. alicea Stoneham 

var. lugari van Someren 
d. teita van Someren 
d. proximans Joicey & Talbot 
d. stevensoni van Someren 
d. entabeni van Someren 
d. moerens Jordan 

d. cinadon Hewitson 

27. phraortes Doubleday [IX : 473] 

28. andranodorus Mabille [IX : 474] 

29. eudoxus (Drury) [VI : 238, X : no] 

e. eudoxus (Drury) 

e. mechowi Rothschild 
e. theresae Le Cerf 
e. hater ae Carpenter 
e. cabacus Jordan 
e. amaurus Poulton 
f. nzoia van Someren 



REVISIONAL NOTES ON AFRICAN CHARAXES 



"3 



e. lucyae van Someren 
e. zambiae van Someren 

30. richelmanni Rober [VI : 236] 
tiridates-group 

31. violetta Grose-Smith [III : 50] 

v. violetta Grose-Smith 

melloni Fox 
v. maritima van Someren 
v. meru van Someren 

32. numenes (Hewitson) [VIII : 241] 

n. numenes (Hewitson) 
n. intermediate cline 

f. laticatena Le Cerf 
n. aequatorialis van Someren 

f. obsolescens Stoneham 
n. neumanni Rothschild 

33. fuscus Plantrou [VIII : 240] 

34. tiridates (Cramer) [VIII : 235] 

t. tiridates (Cramer) 
marica (Fabricius) 

t. intermediate cline 
var. tristis Schultze 
var. angusticandatus Rober 
var. purpurina Rousseau-Decelle 
var. subcaerulea Storace 

t. tiridatinus Rober 
ab. conjuncta Storace 

t. marginatus Rothschild 

35. bipunctatus Rothschild [VIII : 246] 

b. bipunctatus Rothschild 

johnsoni Rousseau-Decelle 
b. intermediate cline 
b. ugandensis van Someren 

36. mixtus Rothschild [VIII : 250] 

37. bubastis Schultze [VIII : 251] 

38. albimaculatus van Someren [VIII : 
252] 

39. barnsi Joicey & Talbot [VIII : 253] 

40. bohemani Felder [VII : 208] 

41. schoutedeni Ghesquiere [II : 220, V : 

79] 

42. montieri Staudinger (VII : 200] 

43. overlaeti Schouteden [Not dealt with 
in text but of doubtful affinity.] 

44. smaragdalis Butler [II : 207] 

s. smaragdalis Butler 
f. beni van Someren 
s. butleri Rothschild 
s. leopoldi Ghesquiere 
s. metu van Someren 
s. caerulea Carpenter & Jackson 
s. toro van Someren 
s. kagera van Someren 
* See p. 127 for cithaeron Felder, inadvertently 



s. elgonae van Someren 
s. homonymus Bryk 

orientalis Joicey & Talbot 
s. kigoma van Someren 
45. xiphares (Cramer) [II : 188, V : 82 
VIII : 259] 

x. xiphares (Cramer) 

f. occidentalis van Son 
x. thyestes (Stoll) 

reducta Rothschild 

elatias Jordan 
x. penningtoni van Son 

^ f. luminosa van Son 
x. draconis Jordan 

$ f. Candida van Son 
x. kenwayi Poulton 

$ f. lutea van Son 
*. bavenda van Son 

9 f. ochreomacula van Son 

$ f. cyanescens van Son 
x. vumbui van Son 
x. woodi van Someren 
x. brevicaudatus Schultze 
*. burgessi van Son 
x. maudei Joicey & Talbot 
*. kulal van Someren 
x. desmondi van Someren 
x. wernickei Joicey & Talbot 
x. kilimensis van Someren 
x. ludovici Rousseau-Decelle [X : 109] 
46*. nandina Rothschild & Jordan [II : 203] 

47. imperialis Butler [VIII : 223] 

1. imperialis Butler 
i. albipunctus Joicey & Talbot 
1. paulianus Rousseau-Decelle 
1. ugandicus van Someren 

f. caerulipunctus van Someren 
i. lisomboensis van Someren 

[ludovici Rousseau-Decelle sensu 

van Someren] 

48. ameliae Doumet [VIII : 228] 

a. ameliae Doumet 
regius Aurivillius 
a. victoriae van Someren 
a. amelina Joicey & Talbot 

49. pythodoris Hewitson [I : 222, X : 105] 

p. pythodoris Hewitson 
p. occidens van Someren 
p. nesaea Grose-Smith 
p. pallida Carpenter 
p. davidi Plantrou 
had rianus-grou p 

50. hadrianus Ward [VIII : 221] 
omitted from this Synoptic List. 



II 4 



V. G. L. van SOMEREN 



h. hadrianus Ward 

dux Staudinger & Schatz 
gabonica Crowley 

h. lecerfi Lathy 
no bilis-group 

51. nobilis Druce [IX : 433] 

n. nobilis Druce 

agabo Distant 

homerus Staudinger 
n. rosemariae Rousseau-Decelle 
n. claudei Le Moult 

52. superbus Schultze [IX : 436] 

53. acraeoides Druce [IX : 429] 

54. fournierae Le Cerf 

f.fournierae Le Cerf 
/. kigeziensis Howarth 

55. lydiae Holland [IX : 481] 
zoo/ina-group 

56. kahldeni Homeyer & Dewitz [IX : 450] 

apicalis Rober 

f. homeyeri Homeyer & Dewitz 

f. bellus Niepelt 

57. zoolina (Westwood) [IX : 444] 

z. zoolina (Westwood) 

f. neanthes (Hewitson) 

f. homochrous Le Cerf 

f. obscuratus Suffert 
z. mqfugensis Jackson 
z. ehmckei Homeyer & Dewitz 

f. phanera Jordan 
z. betsimisaraka Lucas 

relatus Butler 

firmus Le Cerf 

f . beianimena Lucas 

andriba Ward 

freyi (Branczik) 

lambertoni Lathy 
cupa/e-group 

58. eupale (Drury) [IX : 438] 
e. eupale (Drury) 

amasia (Fabricius) 
e. latimargo Joicey & Talbot 

schultzi Rober 

59. subornatus Schultze [IX : 440] 

s. subornatus Schultze 
s. minor Joicey & Talbot 

60. dilutus Rothschild [IX : 441] 

d. dilutus Rothschild 
d. miotoni van Someren 
ngonga van Someren 

61. montis Jackson [IX : 442] 
jahlusa-group 

62. jahlusa (Trimen) [IX : 418] 

j.jahlusa (Trimen) 



/. argynnides Westwood 

j. kigomaensis van Someren 
kigoma van Someren 

/. kenyensis Joicey & Talbot 
f. pallene van Someren 
f . transitional to ganalensis 

j. ganalensis Carpenter 
p/etone-group 

63. pleione (Godart) [IX : 427] 

p. pleione (Godart) 

lichas (Doubleday) 

f. othello Suffert 

f . pallida Lathy 
p. bebra Rothschild 

64. paphianus Ward [IX : 425] 

p. paphianus Ward 
falcata (Butler) 
hamulosa (Weymer) 
p. subpallida Joicey & Talbot 
zingha-group 

65. zingha (Stoll) [IX : 466] 

berenice (Drury) 
etesipe-group 

66. etesipe (Godart) [III : 59] 

e. etesipe (Godart) 
etheta (Godart) 
$ f. castoroides Poulton 
$ f. caeruleotincta Carpenter 

e. abyssinicus Rothschild 

e. patrizii Storace 

e. tavetensis Rothschild 

e. gordoni van Someren 

e. pemba van Someren 

67. penricei Rothschild [III : 65, V : 80] 

p. penricei Rothschild 

ab. peculiaris Lathy 

ab. flavus Lathy 
p. dealbata Joicey & Talbot 
p. tanganyikae van Someren 

$ f. caerulescens van Someren 

68. cacuthis Hewitson [III : 68] 

69. paradoxa Lathy [III : 67] 

70. achaemenes Felder [VI : 207] 

a. achaemenes Felder 

f . fasciatus Suffert 
a. monticola Joicey & Talbot 

f. erythraea Storace 
a. cline monticola x atlantica 
a. atlantica van Someren 

jocaste Butler 
etheocles-gxoup 

71. anticlea (Drury) [VII : 209] 

a. anticlea (Drury) 
horatius (Fabricius) 



REVISIONAL NOTES ON AFRICAN CHARAXES 



"5 



a. proadusta van Someren 
a. cline proadusta x adust a 
a. adusta Rothschild 

a. suna van Someren 

reducta van Someren 

72. baumanni Rogenhofer [VII : 217, 
IX : 482] 

b. baumanni Rogenhofer 
b. tenuis van Someren 

b. interposita van Someren 
b. bwamba van Someren 
b. didingensis van Someren 
b. whytei Butler 
selous Trimen 
b. bamptoni van Someren 

73. opinatus Heron [VI : 202] 

74. thysi Capronnier [VIII : 219] 

75. hildebrandti Dewitz [VIII : 218] [X : 

105] 

h. hildebrandti Dewitz 

talagugae Holland 

galba Distant 
h. gillesi Plantrou 
h. katangensis Talbot 

76. blanda Rothschild [VI : 205] 

b. blanda Rothschild 
b. kenyae Poulton 

77. kheili Staudinger [V : 94] 

78. northcotti Rothschild [V : 96] 

79. guderiana (Dewitz) [VI : 199] 

g. guderiana (Dewitz) 

tanganika Robbe 
g. rabaiensis Poulton 

80. pembanus Jordan [III : 70] 

81. usambarae van Someren & Jackson 
[HI : 73] 

f. collinsi van Someren 

82. contrarius Weymer [V : 119] 

subargentea van Someren & Rogers 
f. conjugens van Someren 

83. peter si van Someren [V : 121] 

84. marieps van Someren & Jackson 

[III : 80] 

85. kar kloof van Someren [III : 81] 

k. karkloof van Someren 
k. capensis van Someren 
86. martini van Someren [III : 96, IX : 

483] 

87. gallagheri van Son [III : 94] 

88. alpinus van Someren & Jackson [III : 

84] 

89. nyikensis van Someren [X : 107] 

90. maccleeryi van Someren [VIII : 257] 



91. grahamei van Someren [V : 115] 

$ f. lacteata van Someren 

92. aubyni van Someren & Jackson [III : 

77] 
a. aubyni van Someren & Jackson 
a. ecketti van Someren & Jackson 

a. australis van Someren & Jackson 

93. chepalungu van Someren [V : 90] 

94. virilis Rothschild [V : 92] 

lenis Jordan 

95. fulgurata Aurivillius [III : 92] 

$ f. lunigera Rothschild & Jordan 
$ f. mima Riley 

96. berkeleyi van Someren & Jackson 

[V : 80] 

b. berkeleyi van Someren & Jackson 

$ f. ngonga van Someren 
b. masaba van Someren 

97. bailey i van Someren [V : 122] 

$ f . pseudocarpenteri van Someren 

98. manica Trimen [III : 86, VIII : 255] 

m. manica Trimen 
m. subrubidus van Someren 
°. f. atribasis van Someren 
$ f. aubergeri van Someren 
$ f. pseudosmaragdalis van Som- 
eren & Jackson 

99. pseudophaeus van Someren [X : 107] 

100. chintechi van Someren [X : 107] 

101. protomanica van Someren [X : 107] 

102. ethalion (Boisduval) [IV : 281] 

Group 1. 
e. ethalion (Boisduval) 
§ f. ethalion (Boisduval) 
$ f. swynnertoni Poulton 
$ f. rosae Butler 
$ f . aurantimacula van Someren 
Group 2. See No. 103 pondoensis 
Group 3. 
e. nyasicus van Someren 
nyasana van Someren 
$ f. nyasicus van Someren 
nyasana van Someren 
$ f. swynnertoni pattern 
$ f. cithaeronoides van Someren 
$ f. suppressa van Someren 
$ f. demaculata van Someren 
$ f. imitans van Someren 
Group 4. 
e. nyanzae van Someren 
Region 1 

$ f. ethalion pattern 
$ f. howardi van Someren & 
Jackson 



Ii6 



V. G. L. van SOMEREN 



Region 2 

$ f. ethalion pattern 
Region 3 

°. f. ethalion pattern 
°. f. rosae pattern 
Region 4 

$ f. ethalion pattern 
Group 5. 
e. littoralis van Someren 
Region 1 

°. f. ethalion pattern 
§ f. rosae pattern 
Region 2 

$ f. ethalion pattern 
<j> f. rosae pattern 
$ f. swynnertoni pattern 
e. intergrading cline between Groups 
3 and 5 
$ f . ethalion pattern 
$ f. rosae pattern 
°. f. swynnertoni pattern 
Group 6. 
e. kikuyuensis van Someren 
$ f . ethalion pattern 
$ f. rosae pattern 
$ f. swynnertoni pattern 
Group 7. 
c. marsabitensis van Someren 
$ f. swynnertoni pattern 

103. pondoensis van Someren [X : 108] 

104. etheocles (Cramer) [V : 90] 
Region 1 

e. etheocles (Cramer) 

$ f. etheocles (Cramer) 

$ f. alladinis Butler 

$ f . fulgens Rothschild 

<j> f. regalis Rothschild 

o* f. ephyra Godart 
hollandi Butler 

<$ f. carteri Butler 

<J f. catochrous Staudinger 
Region 2 
e. biinclinata van Someren 

$ f . etheocles pattern (two vars.] 

$ f. ochracea pattern 

9 f. alladinis pattern 

$ f . regalis pattern 

<J f. ephyra pattern 

cT f . carteri pattern 

<$ f. catochrous pattern 
Region 3 
c. ochracea van Someren & Jackson 

$ f. ochracea Rothschild 



$ f. ochreata van Someren & 
Jackson (= alladinis X regalis) 
$ f. seriata Rothschild 
(J f. violacea Rothschild 
Region 4 
e. carpenteri van Someren & Jack- 
son 
f. carpenteri Poulton 
$ f. carpenteri Poulton and vars. 
°. f . pallidimacula van Someren & 

Jackson and vars. 
o* f. carpenteri van Someren & 

Jackson 
<J f. near carteri Butler 
(J f. near catochrous Staudinger 
Region 5 
e. evansi van Someren & Jackson 
f. evansi van Someren & Rogers 
$ f. evansi van Someren & Jack- 
son and vars. 
$ f. conjuncta van Someren & 

Jackson 
<J f. evansi van Someren & Jackson 
(near carteri) and vars. 
105. t'l'oZa Butler [V : 125] 
v. Wo/a Butler 
v. picta van Someren & Jackson 

$ f. vansomereni Poulton 
v. suk Carpenter & Jackson 

$ f. kirkoides Carpenter & Jackson 
$ f. achaemenesopsis Carpenter & 

Jackson 
$ f . intermedia Carpenter & Jackson 
$ f. albifascia Poulton 
v. daria Rothschild 
v. chanleri Holland 
v. kirki Butler 
? f. Airfo Butler 
$ f. albifascia Poulton 
$ f. rogersi Poulton 
$ f. handari Poulton 
v. diversiforma van Someren & 
Jackson 
$ f. diversiforma van Someren & 

Jackson 
$ f. purpurea van Someren & 

Jackson 
$ f. viridicaerulea van Someren & 

Jackson 
$ f . caerulescens van Someren & 

Jackson 
$ f. albocaerulea van Someren & 
Jackson 



REVISIONAL NOTES ON AFRICAN CHARAXES 



117 



$ f. albimacula van Someren & 

Jackson 
$ f. ochremaculata van Someren & 

Jackson 
$ f. cupreopurpurea van Someren 

& Jackson 
v.fagini Storace 

106. phaeus Hewitson [X : 108] 

$ f . phaeus Hewitson 
$ f. corydoni Rothschild 

107. vansoni van Someren [X : 108] 

$ f. vansoni van Someren & Jack- 
son 

108. variata van Someren [X : 108] 

$ f. variata van Someren 
9 f . tricolor van Someren 
$ f. rosella van Someren 
$ f. cottrelli van Someren 

109. loandae van Someren [X : 108] 

$ f. loandae van Someren 
$ f. primitiva van Someren 
? f. basiviridis van Someren 
$ f. violitincta van Someren 
$ f. vansonoides van Someren 
$ f . protokirki van Someren 
$ f. instabilis van Someren 
no. brainei van Son [X : 108] 
in. cedreatis Hewitson [V : 85] 
lutacea Rothschild 
9 f. cedreatis Hewitson 
$ f. protocedreatis Poulton 



$ f. inexpectata van Someren 
$ f. vetula Rothschild 
$ f. pseudosmaragdalis van Som- 
eren & Jackson 
$ f. dewitzi Butler 

112. mafuga van Someren [V : 97, X : 106] 
nichetes-group 

113. nichetes Grose-Smith [IX : 463] 

n. nichetes Grose-Smith 

hamatus Dewitz 

ogovensis Holland 
n. leoninus Butler 
n. pantherinus Rousseau-Decelle 
laodice-group 

114. laodice (Drury) [IX : 460] 

nesiope (Hewitson) 
lycurgus (Fabricius) 

115. zelica Butler [IX : 457] 

z. zelica Butler 

z. depuncta Joicey & Talbot 

z. toyoshimai Carcasson 

116. porthos Grose-Smith [IX : 454] 

p. porthos Grose-Smith 

tnidas Staudinger 
p. katangae Rousseau-Decelle 
p. dummeri Joicey & Talbot 
p. gallayi van Someren 

117. dunkeli Rober [IX : 461] 

118. doubledayi Aurivillius [IX : 454] 

119. mycerina (Godart) [IX : 452] 

nausicaa Staudinger 



EUXANTHINAE Rydon 

EUXANTHE Htibner 
Subgenus EUXANTHE Hiibner 



1. eurinome (Cramer) [X: 79] 

e. eurinome (Cramer) 

$ f . johnsoni Howarth 
e. ansellica Butler 

f. burgeoni Le Cerf 

var. radiata van Someren & Rogers 
c. celadon Le Cerf 
e. elgonae van Someren 
e. birbirica Ungemach 

2. crossleyi (Ward) [X : 83] 

c. crossleyi (Ward) 
f. niepelti Bryk 



3- 



c. ansorgei Rothschild & Jordan 

f. babbingtoni Stoneham 
c. magnifica Rebel 

intermedia Joicey 
c. claudiae Rousseau-Decelle 
wakefieldi (Ward) [X : 85] 

f. rubiginea Le Cerf 
madagascariensis (Lucas) [X : 86] 
amakosa (Boisduval) 



n8 



V. G. L. van SOMEREN 



Subgenus HYPOMELAENA Aurivillius 



trajanus (Ward) [X : 87] 
t. trajanus (Ward) 

schatzi Staudinger 
t. vansomereni Poulton 
t. gabonicus Le Cerf 
t. antonius Rousseau-Decelle 
t. nigeriae van Someren 



1. publius Staudinger [X : 75] 

p. publius Staudinger 
rectifascia Weymer 
f. rectifascia Weymer 
f. moderata Gaede 
p. centralis van Someren 
p. kigoma van Someren 

2. ussheri (Butler) [X : 70] 

u. ussheri (Butler) 



6. tiberius Grose-Smith [X : 90] 
t. tiberius (Grose-Smith) 

f. tiberiella Strand 
t. meruensis van Someren 



PALLINAE Rydon 

PALLA Hiibner 



$ f. ferruginea Schultze 
$ f. dobelli Hall 
u. interposita Joicey & Talbot 

3. decius (Cramer) [X : 69] 

f. Sagittarius Rousseau-Decelle 

4. violinitens (Crowley) [X : 73] 

v. violinitens (Crowley) 
v. coniger (Butler) cline 
v. bwamba van Someren 



ACKNOWLEDGEMENTS 

In submitting this, the last part of Revisional Notes on African Charaxes, I wish 
to acknowledge with grateful thanks the assistance I have received from individuals 
and members of museums without whose help the work could not have been under- 
taken. Foremost amongst these associates has been Mr T. G. Howarth of the 
Department of Entomology, British Museum (Natural History), who has not only 
provided me with specimens but has also sent me photographs of types deposited 
in the BMNH, Mr C. F. Huggins of the same Department who has mounted the 
photographs for the plates and Mr R. Smiles who has dealt with the maps and 
prepared the bibliography. 

The extensive editing of the series - no light task - has been undertaken by Mr 
Howarth and Mr D. E. Kimmins, and I tender my thanks. I wish to record my 
indebtedness to Dr A. H. B. Rydon, Chailey, Sussex, who has been of great assistance 
with the checking of the typescript and proofs of this final part; to Dr P. Viette 
andDr G. Bernardi, Museum National d'Histoire Naturelle, Paris; to Monsieur L. A. 
Berger, Musee Royal de FAfrique Centrale, Tervuren; Dr H. J. Hannemann, Museum 
fur Naturkunde, Berlin; Dr F. Kasy, Naturhistorisches Museum, Vienna; Mr E. 
Taylor, University Museum, Oxford; the late Dr R. M. Fox, Carnegie Museum, 
Pittsburgh; Dr F. Rindge, American Museum of Natural History, New York; 
the late Dr G. van Son, Transvaal Museum, Pretoria; Dr E. Pinhey, National 
Museum, Bulawayo; Dr C. M. McCleery, Zomba, Malawi; Mr H. T. Martin, Limbe, 
Malawi; the late Mr K. M. Pennington, Balgowan, Natal; Mr C. G. C. Dickson, 
Cape Town, South Africa; Mr Lawson, Durban Museum, Natal; Dr R. H. Carcasson, 
late of the National Museum, Nairobi; Mr M. P. Clifton, National Museum, 
Nairobi; Major I. Grahame, Lamarsh, Suffolk; Mr H. D. van Someren, Kitale, 



REVISIONAL NOTES ON AFRICAN CHARAXES 119 

Kenya; Monsieur Jacques Plantrou, Paris; Prof. Schumattera, Zoological Depart- 
ment, Nairobi University. 

My special thanks are due to Mr D. G. Sevastopulo of Mombasa for the descriptions 
of the early stages of Euxanthe tiberius tiberitis Grose-Smith, together with photo- 
graphs illustrating the various stages. 

To all those who have assisted me with material and information during the 
prolonged period of preparation of these revisional notes, I renew my thanks. 



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79 (A) 8 : 3. 

1914- Neue Charaxiden aus dem tropischen Afrika. Ent. Rdsch. 31 : 82. 

1916. Die Charaxiden und Apaturiden der Kolonie Kamerun. Arch. Biontol. 4 : 81-129, 

6 pis, 1 map. 
191 7. Ergebnisse 2. Deutschen Zentral- Afrika Expedition 1910-1911. Lepidoptera. 

Ergebn. Zw. dt. zent. Afr. Exped. 1 (12) : 511-597, 7 pis. 
Seba, A. 1765. Locupletissimi rerum naturalium thesauri 4. 226 pp., 108 pis. Amsterdam. 



REVISIONAL NOTES ON AFRICAN CHARAXES 125 

Sharpe, E. M. 1904. On new species of butterflies from equatorial Africa. Entomologist 

37 : 131-134. 
Staudinger, O. 1891. Neue exotische Lepidopteren. Dt. ent. Z. Iris 4 : 61-157. 

1892. Ueber die Arten der Palla Decius-Gruppe. Dt. ent. Z. Iris 5 : 264-268. 

Staudinger, O. & Schatz, E. 1884-88. Exotische Schmetterlinge, 1, pp. 1-333, 100 pis. Fiirth. 

1896. Neue exotische Tagfalter. Dt. ent. Z. Iris 9 : 193-240. 

Stichel, H. 1939. In Bryk, F., Lepid. Cat. 91 : 375-542. 

Stoneham, H. F. 193 i. A new species of Charaxes, from Kenya Colony. Bull. Stonekam 

Mus. no. 5 : [1-2]. 
1931. A new form of Charaxes castor, Cram., from Kenya Colony. Bull. Stoneham 

Mus. no. 6, pp. 1. 
1931. A New Form of Charaxes numenes, Hew., from Kenya Colony. Bull. Stoneham 

Mus. no. 7 : [1]. 
1932. Two new races of Charaxes pollux Cr., from Kenya, British East Africa. Bull. 

Stoneham Mus. no. 9 : 1-2. 
1936. Notes on Charaxes, O., from Kenya and Uganda, Brit. E. Africa. Bull. Stoneham 

Mus. no. 29 pp. 3. 
1943. New forms of Lepidoptcra of the subfamily Charaxidinae, from East Africa. 

Bull. Stoneham Mus. no. 46, 3 pp. 
1951-65. Butterflies of western Kenya. Pts 1-11, pp. 1-80. Stoneham Museum 

Publications, Nairobi. 
Storace, L. 1948. Descrizione di nuove forme africane del gen. Charaxes O. (Lepidoptera, 

Nymphalidae). Annali Mus. civ. Stor. nat. Giacomo Doria 63 : 132-141. 
1949. Spedizione zoologica del Marchese Saverio Patrizi nel Basso Giuba e nell'Oltregiuba, 

VI-VIII. 1934. Lepidotteri diurni. Annali Mus. civ. Stor. nat. Giacomo Doria 64 : 12-29. 
Stoll, C. 1790. In Cramer, P., Uitlandsche Kapellen 1—4, Suppl. Amsterdam & Utrecht. 
Strand, E. 191 i. Die in Berliner Museum vorhandenen Ludia und Holocera Arten, nebst 

Beschreibungen weiterer van Prof. Dr. J. Vosseler gesammelter afrikanischer Gross- 

schmetterlinge. Dt. ent. Z. Iris. 25 : 110-121. 
Suffert, E. 1904. Neue Nymphaliden aus Africa. Dt. ent. Z. 17 : 108-123. 
Talbot, G. 1921. New Lepidoptera collected by Mr T. A. Barns in East Central Africa. 

Bull. Hill Mus. Witley 1 : 40-166, 14 pis. 
1927. Charaxes alticola Griinb. §, and remarks on Charaxes boueti Feisth. (Lep. Nympha- 
lidae). Entomologist 60 : 109-110. 
1928. New forms of African Nymphalidae (Lep. Rhop.) in the Musee du Congo, Tervuren. 

Bull. Hill Mus. Witley 2 : 229-231. 

1929. A note on Charaxes boueti centralis Neustt. (1929). Bull. Hill Mus. Witley 3 : 148. 

1932. Four new forms of Rhopalocera. Bull. Hill Mus. Witley 4 : 288-289. 

1932. A note on the habits of some Charaxes observed on Mount Kenya and the descrip- 
tions of a new race of Charaxes. Bull. Hill Mus. Witley 4 : 181. 
1932. The female of Charaxes alticola Griinb., associated with the female of C. ansovgei 

ruandana Talb., as a mimic of C. brutus Cram. &c. Proc. ent. Soc. Lond. 7 : 9. 
Trimen, R. 1862-1866. Rhopalocera Africae Australis. 353 pp., 7 pis. London and Cape 

Town. 
1894. On a collection of butterflies made in Manica, tropical south-east Africa by Mr 

F. C. Selous, in the year 1892. Proc. zool. Soc. Lond. 1894 : 14-82, pis 4-6. 
Trimen, R. & Bowker, J. H. 1887. South African butterflies 1 : 348 pp. London. 
Thurau, F. 1903. Neue Rhopaloceren aus Ost Afrika. Ergebnisse der Nyassa-See-und 

King-Gebirgs-Expedition der Hermann und Elise geb. Heckmann-Wentzel-Stiftung. 

Berl. ent. Z. 48 : 11 7-143, 1 pi. 
Ungemach, H. 1932. Contribution a l'etude des Lepidopteres d'Abyssinie. (I re partie, 

Rhopaloceres) . Mem. Soc. Sci. nat. phys. Maroc 32 : 1-122, 2 pis. 
van Someren, V. G. L. 1932. Butterflies of Kenya and Uganda [Suppl. i] JIE. Africa Uganda 

nat. Hist. Soc. nos. 42-43 (1931) : 141-172. 



126 V. G. L. van SOMEREN 

van Someren, V. G. L. 1935. Ibidem. [Suppl. 2] Jl E. Africa Uganda nat. Hist. Soc. 12 : 
147-199. 

1939. Ibidem. [Suppl. 3] // E. Africa Uganda nat. Hist. Soc. 14 : 15-100. 

1939. New and little-known Lepidoptera from Kenya and Uganda. Jl E.Africa Uganda 

nat. Hist. Soc. 14 : 172-180. 
1957. Two apparently undescribed female Charaxes (Lepidoptera : Nymphalidae). 

Proc. R. ent. Soc. Lond. (B) 26 : 177. 
1958. A new species and a new female form of "Charaxes" from Kenya Colony. Novos 

Taxa ent. 11 : 1-12, 2 pis. 
1962. A new Charaxes (Lepidoptera: Nymphalidae) from the northern frontier province 

of Kenya. Proc. R. ent. Soc. Lond. (B) 31 : 44-66, pis 1—2. 
1963. Revisional notes on African Charaxes. Part I. Bull. Br. Mus. nat. Hist. (Ent.) 

13 : 198-242, 5 maps, 19 plates. 

1964. Ibidem. Part II. Bull. Br. Mus. nat. Hist. (Ent.) 15 : 181-235, 4 maps, 23 pis. 

1966. Ibidem. Part III. Bull. Br. Mus. nat. Hist. (Ent.) 18 : 45-101, 5 maps, 16 pis. 

1966. Ibidem. Part IV. Bull. Br. Mus. nat. Hist. (Ent.) 18 : 277-316, 4 maps, 9 pis. 

■ 1967. Charaxes jocaste Butler, 1865 (Insecta, Lepidoptera): proposed suppression under 

the Plenary Powers, Z.N.(S.) 1806. Bull. zool. Nom. 24 : 255-256. 
1968. Charaxes porthos Grose-Smith. A brief review of its distribution and subspeciation. 

Bull. Inst. fr. Afr. noire 30 : 221-225, 9 figs. 
1969. Revisional Notes on African Charaxes (Lepidoptera : Nymphalidae). Part V. 

Bull. Br. Mus. nat. Hist. (Ent.) 23 : 75-166, 29 pis, 8 maps, 31 text-figs. 

■ 1970. Ibidem. Part VI. Bull. Br. Mus. nat. Hist. (Ent.) 25 : 197-49, 6 maps, 11 pis. 

1971. Ibidem. Part VII. Bull. Br. Mus. nat. Hist. (Ent.) 26 : 181-226, 11 pis, 6 maps. 

1972. Ibidem. Part VIII. Bull. Br. Mus. nat. Hist. (Ent.) 27 : 215-264, 7 maps, 12 pis. 

1974- Ibidem. Part IX. Bull. Br. Mus. nat. Hist. (Ent.) 29 : 415-487, 9 maps, 18 pis. 

van Someren, V. G. L. & Jackson, T. H. E. 1952. The Charaxes etheocles-ethalion complex: 

a tentative reclassification of the group (Lepidoptera: Nymphalidae). Trans. R. ent. 

Soc. Lond. 103 : 257-284. 
1957- The Charaxes etheocles-ethalion complex (Lepidoptera : Nymphalidae) Supplement 

No. 1. Ann. Transv. Mus. 23 : 41-58, 1 map, pis 3-4. 
van Someren, V. G. L. & Rogers, K. St.A. 1928a. The butterflies of Kenya & Uganda. 

Part 6. Jl E. Africa Uganda nat. Hist. Soc. 30 (1927) : 57-69, 6 pis. 

■ 19286. Ibidem. Part 7. // E. Africa Uganda nat. Hist. Soc. 31-32 : 111-158; 29 pis. 

1929. Ibidem. Part 8. Jl E. Africa Uganda nat. Hist. Soc. 33-34 (1928) : 3-54. 

1931. Ibidem. Part 9. // E. Africa Uganda nat. Hist. Soc. 38-39 (1930) : 18-33. 

1932. Butterflies of Kenya and Uganda. //. E. Africa Uganda nat. Hist. Soc. 42-43 

(1931) : 141-172, pis 108-121. 
van Someren, V. G. L. & van Someren, R. A. L. 1926. The life-histories of certain African 

Nymphalid butterflies of the genera Charaxes, Palla and Euxanthe. Trans, ent. Soc. 

Lond. 74 : 333-354, pis 74-80. 
van Son, G. 1935. New butterflies from South Africa. Ann. Transv. Mus. 15 : 485-489. 

1936. Description of a new race of Charaxes xiphares (Cr.) from Southern Rhodesia, 

(Lepid.) Proc. R. ent. Soc. Lond. (B) 5 : 201-206, pi. 2. 

- 1953- A revision of the subspecies and forms of Charaxes cithaeron Felder and Charaxes 

xiphares (Cr.). Ann. Transv. Mus. 22 : 219-230, 4 pis. 

1962. New African Butterflies, 1. /. ent. Soc. sth. Afr. 25 : 140-148, pi. 2. 

■ 1963. Ibidem, 2. Novos Taxa ent. no. 29 : 1-11, 1 pi. 

1966. A new subspecies of Charaxes viola Butler from South-West Africa. Novos 

Taxa ent. no. 49 : 3—7. 
Wallengren, H. D. J. 1857. Kafferlandets Dag-Fjarilar. K. svenska VetenskAkad. Handl 

2 : i-55- 
Ward, C. 1871. Descriptions of new species of African diurnal Lepidoptera. Entomologist's 
mon. Mag. 8 : 34-36, 11 8-1 22. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



127 



Ward, C. 1872. Descriptions of new species of diurnal Lepidoptera from Madagascar. 

Entomologist' s mon. Mag. 9 : 2-3. 
1873. Descriptions of new species of African Lepidoptera. Entomologist's mon. Mag. 

9 : 147-148, 151-152, 209-210. 
Watkins, H. T. G. 1923. Notes on the butterflies of the Banks Collection. Entomologist 

56 : 204-209. 
Westwood, J. O. 1864. Charaxes Argynnides, Westw. Proc. ent. Soc. Lond. (3) 2 : 10. 

1874. Thesaurus entomologicus Oxoniensis, 205 pp., 40 pis. Oxford. 

Westwood, J. O., Doubleday, E. & Hewitson, W. C. 1850-52. The genera of diurnal 

Lepidoptera 2 : 534 pp., 81 pis. London. 
Weymer, G. 1892. Exotische Lepidopteren VI. Stettin, ent. Ztg 53 : 79-125. 
Wilson, C. E. 1950. Butterflies of the northern and central Sudan. Mem. Res. Div. Min. 

Agr. Sudan no. 12 : 222-224. 



APPENDIX 



The following entry was inadvertently omitted from p. 113 of the Synoptic 
List. 



45a cithaeron Felder [II : 225, X 
c. cithaeron Felder 

ab. whitei van Someren 
c. joanae van Someren 



105] 



c. nyasae van Someren 

ab. griseus Schultze 
c. kennethi Poulton 
c. nairobicus van Son 



INDEX TO PARTS I-X 

This is a complete index to the specific and infraspecific names in all ten parts of the 
series. The Roman numbers refer to the parts. Synonyms are in italics. 



abyssinicus, III : 61, 
achaemenes, VI : 207, X : 114 
achaemenesopsis, V : 132 
acraeoides, IX : 429, X : 114 
acuminatus, I : 211, V : 77, X : 111 
adusta, VII : 212 
aequatorialis, VIII : 244 
aequidistans , VII : 206 
aeson, VII : 202 
agabo, IX : 433 
aginga, VII : 186 
albifascia, V : 131, 133 
albimacula, V : 143 
albimaculatus, VIII : 252, X : 113 
albipunctus, VIII : 225 
albocaerulea, V : 143 
albofascia, cynthia, V : 152 
albofascia, lucretius, VII : 196 
alcyone, VI : 219 
alicea, I : 236 
alladinis Butler, V : 100 
alladinis Dewitz, V : 90 
alpinus, III : 84, X : 107, 115 
alticola, VI : 230 
amakosa, X : 86 



amasia, IX : 438 

amaurus, VI : 245 

ameliae, VIII : 228, X : 113 

analava, IX : 472, X : 111 

andara, VI : 221, X : 112 

andranodorus, IX : 474, X : 112 

andriba, IX : 449 

angusticaudatus, VIII : 240 

angusticlavius, V : 151 

angustus, VI : 216 

ansellica, X : 81 

ansorgei (Charaxes), IV : 303, X : 104, 112 

ansorgei (Euxanthe), X : 84 

antamboulou, IX : 472, X : 111 

anticlea, VII : 209, X : 108, 114 

antiqua, VII : 185 

antiquus, VI : 220 

antonius, X : 89 

apicalis, IX : 452 

arabica, VII : 193 

argynnides, IX : 420 

arthuri, VII : 188 

atlantica, VI : 211 

atribasis, VIII : 255 

aubergeri, VIII : 256 



128 



V. G. L. van SOMEREN 



aubyni, III : 77, X : 115 
aurantiaca, V : 157 
aurantimacula, IV : 283 
australis, III : 79 
austrinus, IX : 476 
azota, VII : 206 

babbingtoni (Euxanthe), X : 84 

babingtoni (Charaxes), VII : 197 

baileyi, V : 122, X : 115 

balfouri, X : 111 

bamptoni, IX : 482 

baringana, VII : 191 

barnsi (barnesi), VIII : 253, X : 113 

basiviridis, V : 145 

baumanni, VII : 216, IX : 482, X : 115 

bavenda, II : 193 

bebra, IX : 428 

bellus, IX : 452 

berenice, IX : 466 

berkeleyi, III : 74, V : 80, X : 115 

bertrami, IX : 479 

betanimena, IX : 449 

betsimisaraka, IX : 448 

biiclinata, V : 102 

bipunctatus, VIII : 246, X : 113 

birbirica, X : 82 

blanda, VI : 205, X : 115 

bohemani, VII : 208, X : 113 

boueti, VI : 223, X : 111 

brachycauda, IX : 478 

brainei, V : 147, X : 108, 117 

brevicaudatus, II : 195 

brunnescens, I : 205 

brunneus, VIII : 235 

brutus, VI : 214, X : 112 

bubastis, VIII : 251, X : 113 

bungense, IV : 312 

burgeoni, X : 93 

burgessi, II : 197 

busogus, VIII : 234 

butleri, II : 212 

bwamba (Charaxes), VII : 221 

bwamba (Palla), X : 74 

cabacus, VI : 244 

cacuthis, III : 68, X : 114 

caerulea, II : 213 

caerulescens van Someren, V : 80 

caerulescens van Someren & Jackson, V : 143 

caerulipunctus, VIII : 227 

cajus, VI : 214 

caliginosa, VII : 196 

calliclea, VII : 206 



cameroonensis, V : 151 

Candida, II : 192 

candiope, IX : 467, X : in 

capensis, III : 83 

carpenteri, V : 108 

carteri, V : 85, 102 

castor, VII : 183, X : 112 

catenaria, VII : 205 

catochrous, V : 102, 111 

cedreatis, V : 85, X : 117 

celeddn, X : 81 

centralis (Charaxes), VI : 232 

centralis (Palla), X : 77 

chanleri, V : 133 

Charaxes, X : in 

Charaxinae, X : 111 

chepalungu, V : 90, X : 115 

chintechi, III : 90, X : 107, 115 

cinadon Butler, IX : 474 

cinadon Hewitson, I : 231 

cithaeron, II : 225, X : 105, 127 

cithaeronoides, IV : 287 

cizeyi, V : 152 

claudei, IX : 435 

claudiae, X : 85 

collinsi, X : 105 

comoranus, VII : 189 

coniger, X : 74 

conjugens, V : 121 

conjuncta Storace, VIII : 240 

conjuncta van Someren & Jackson, V : 113 

contrarius, V : 119, X : 115 

coryndoni, V : 137 

cottrelli, acuminatus subsp., I : 213 

cottrelli, viola variata $, V : 140 

cowani, IX : 471, X : 111 

crossleyi, X : 83, 117 

cryanae, I : 235 

cupreopurpurea, V : 144 

cyanescens, II : 194 

cynthia, V : 150, X : in 

cytila, III : 86 

daria, V : 134 

davidi, X : 105 

dealbata, III : 66 

decius, X : 69, 118 

defulvata, IX : 480 

demaculata, IV : 288 

depuncta Joicey & Talbot, IX : 458 

depuncta Le Cerf, X : 89 

desmondi, II : 201, VIII : 262 

dewitzi, V : 90 

didingensis, VII : 221 



INDEX 



129 



dilutus, IX : 441, X : 109, 114 

diversiforma, V : 141, 143 

dobelli, X : 71 

doubledayi, IX : 454, X : 117 

draconis, II : 191 

druceanus, I : 228, V : 78, X : 112 

dummeri, IX : 456 

dunkeli, IX : 461, X : 117 

dux, VIII : 221 

ecketti, III : 78 

ehmckei, IX : 448 

elatias, II : 189 

elgonae (Charaxes), II : 215 

elgonae (Euxanthe), X : 82 

entabeni, I : 233 

ephyra, V : 101 

epijasius, I : 201 

erythraea, VI : 210 

etesipe, III : 59, X : 114 

ethalion, IV : 281, X : 108, 109, 115 

etheocles, V : 99, X : 116 

etheta, III : 59 

eudoxus, VI : 239, X : no, 112 

eupale, IX : 438, X : 114 

eurinome, X : 79, 117 

Euxanthe, X : 78 

Euxanthinae, X : 95, 117 

evansi, V : 112 

fagini, V : 149 

falcata, IX : 425 

fallax, VI : 236 

fasciatus, VI : 209 

feisthameli, I : 202 

ferruginea, X : 70 

firmus, IX : 448 

flavescens, VI : 227 

flavifasciatus, VII : 187 

flavimarginalis, VII : 186 

flavus, III : 66 

fournierae, IX : 431, X : 114 

fractifascia, VI : 217 

freyi, IX : 449 

fulgens, V : 101 

fulgurata. III : 92, X : 115 

fulvescens, I : 210, X : 103, in 

fuscus, VIII : 240, X : 113 

gabonica, VIII : 221 
gabonicus, X : 89 
galba, VIII : 218 
gallagheri, III : 94, X : 115 
gallayi, IX : 457 



ganalensis, IX : 422 
gazanus, IV : 314 
geminus, IV : 313 
ghanaensis, VI : 227 
gillesi, X : 105 
gordoni, III : 63 
grahamei, V : 115, X : 115 
griseus, II : 230 
guderiana, VI : 199, X : 115 
guineensis, V : 150 

hadrianus, VIII : 221, X : 113 

hamatus, IX : 463 

hamulosa, IX : 425 

handari, V : 131 

hansali (i), VII : 190, X : 108, 112 

harrisoni, I : 202 

hildebrandti, VIII : 218, X : 105, 115 

hollandi, V : 99 

homerus, IX : 433 

homeyeri, IX : 451 

homonymus, II : 217 

horatianus, VII : 213 

howardi, IV : 293 

Hypomelaena, X : 94, 95, 118 

illuminata, V : 128 

imitans, IV : 290 

imperialis, VIII : 223, X : 109, 113 

inexpectata, V : 88 

instabilis, V : 146 

intermedia (Charaxes), V : 133 

intermedia (Euxanthe), X : 84 

intermedius, VII : 196 

interposita (Charaxes), VII : 220 

interposita (Palla), X : 72 

jacksoni, IV : 304 
jacksonianus, VIII : 234 
jahlusa, IX : 418, X : 108, 114 
jasius, I : 201, X : 112 
joanae, II : 227 
jocaste, VI : 211 
johnsoni, VIII : 246 
johnsoni, X : 80 
Junius, VI : 217 

kagera, II : 218 
kahldeni, IX : 450, X : 114 
karkloof, III : 81, X : 115 
katangae, IX : 455 
katangensis, VIII : 219 
katerae, VI : 243 
kenyae, VI : 206 



130 



V. G. L. van SOMEREN 



kenyensis, IX : 420 

kennethi, II : 231 

kenwayi, II : 192 

kheili, V : 94, X : 115 

kigezia, I : 218 

kigeziensis, IX : 432 

kigoma, jahlusa subsp. IX : 423 

kigoma, smaragdalis subsp., II : 219 

kigoma (Palla), X : 77 

kigomaensis, jahlusa subsp., IX : 423, X : 108 

kikuyuensis, IV : 296 

kilimanjarica, IV : 309 

kilimensis, V : 82, VIII : 260 

kinduana, V : 153 

kirki, V : 130 

kirkoides, V : 132 

kivuanus, I : 235 

kulal, II : 200 

kulalae(nsis),hansali subsp., VII : 192, X :io8 

kulalensis, acuminatus subsp., I : 217 

kungwensis, IV : 306 

lacteata, V : 117 

lactetinctus (lactitinctus), VIII : 232, 

IX : 4 86, X : 112 
lambertoni, IX : 449 
laodice, IX : 460, X : 117 
lasti, VI : 232, X : in 
laticatena, VIII : 243 
laticinctus, I : 205 
latimargo, IX : 440 
lecerfi, VIII : 222 
lemosi, VII : 198 

lenis, V : 92 

leoninus, IX : 465 

leopoldi, II : 212 

levicki, IV : 307 

liberiae, I : 204 

lichas, IX : 427 

lisomboensis, X : 109 

littoralis, IV : 294 

loandae, V : 144, X : 108, 117 

lucida, VII : 195 

lucretius, VII : 194, X : 112 

lucyae, X : 104 

ludovici, imperialis subsp., VIII : 227, 

X : 109 

ludovici, xiphares II subsp., 195, VIII : 262 

X : 109 
lugari, I : 237 
luminosa, II : 191 
lunigera, III : 92 
lutacea, V : 85 
lutea, II : 193 



lycurgus, IX : 460 

lydiae, IX : 481, X : no, 114 

lysianassa, V : 150 

maccleeryi, X : 115 

macclounii, VI : 227 

maculatus, I : 201 

madagascariensis, X : 86, 117 

mafuga, V : 97, X : 106, 117 

mafugensis, IX : 447 

rnagnifica, X : 84 

manica, III : 86, VIII : 255, X : 107, 115 

marginatus, VIII : 239 

marica, VIII : 236 

marieps, III : 80, X : 115 

maritima, III : 52 

marsabitensis, IV : 298 

martini, III : 96, IX : 483, X : 106, 115 

masaba, V : 82 

maua, IV : 316 

maudei, II : 198 

mawamba, V : 153 

maximus, VII : 197 

mccleeryi (maccleeryi), VIII : 257, X : 115 

mechowi, VI : 240 

melas, I : 202 

meru, III : 53 

meruensis, X : 92 

metu, II : 219 

midas, IX : 454 

mima, III : 93 

minor, IX : 441 

miotoni, X : 109 

mixtus, VIII : 249, X : 113 

mlanji, I : 212 

moderata, X : 76 

moerens, I : 232 

monitor, I : 210 

monticola, VI : 209 

montieri, VII : 200, X : 113 

montis, IX : 442, X : 114 

mukuyu, V : 158 

murina, I : 201 

musakensis, X : no 

mycerina, IX : 452, X : 117 



nairobicus, II : 232 
nandina, II : 203, X : 
natalensis, VI : 220 
nausicaa, IX : 452 
neanthes, IX : 446 
nesaea, I : 223 
nesiope, IX : 460 
neumanni, VIII : 425 



113 



INDEX 



131 



ngonga, berkeleyi f., V : 80 

ngonga, dilutus subsp., IX : 442, X : 109 

nichetes, IX : 463, X : 117 

niepelti, X : 83 

nigeriae, X : 89 

nigrescens, VI : 217 

nobilis, IX : 433, X : 114 

northcotti, V : 96, X : 115 

nothodes, VII : 204 

numenes, VIII : 241, X : 113 

nyanzae, IV : 292 

nyasae, II : 229 

nyasana van Someren, IV : 286, X : 109 

nyasana Butler, VII : 206 

nyasicus, X : 109 

nyika, I : 214 

nyikensis, III : 85, X : 107, 115 

nzoia, VI : 246 

obscura, I : 235 

obsolescens, VIII : 244 

obudoensis, V : 77 

occidens, I : 223, V : 79 

occidentalis, II : 189 

ochracea, V : 106 

ochreata, V : 107 

ochrefascia, III : 78 

ochremaculata, V : 143 

ochreomacula, II : 193 

ochretincta, III : 79 

octavus, X : 103, 112 

odysseus, VII : 199, X : 112 

ogovensis, IX : 463 

ongeus, IV : 312 

opinatus, VI : 202, X : 115 

oreas, I : 216 

orientalis Joicey & Talbot, II 1217 

orientalis Lanz, VII : 187 

othello, IX : 427 

overlaeti, X : 113 

pagenstecheri, I : 203 
Palla, X :69, 118 
pallene, IX : 421 
pallida Carpenter, I : 224 
pallida Lathy, IX : 427 
pallidimacula, V : no 
Pallinae, X : 95, 118 
pantherinus, IX : 465 
paphianus, IX : 425, X : 114 
paradoxa, III : 67, X : 114 
parvicaudatus, V : 154 
patergodarti, X : 103 



patrizii, III : 62 

pauliana, VIII : 227 

paulianus, VIII : 227 

peculiaris, III : 65 

pelias, I : 205, X : 112 

pemba. III : 64 

pembanus, III : 70, X : 115 

penningtoni, II : 190 

penricei, III : 65, V : 80, X : 114 

petersi, V : 121, X : 115 

phaeus, V : 136, X : 108, 117 

phanera, IX : 448 

phoebus, VI : 212, X : 112 

phraortes, IX : 473, X : 112 

picta, V : 128 

pleione, IX : 427, X : 114 

pollux, IV : 311, X : 112 

pondoensis, IV : 285, X : 108, 116 

porthos, IX : 454, X : 117 

primitiva, V : 144 

proadusta, VII : 211 

propinqua, V : 155 

protocedreatis, V : 87 

protoclea, VII : 202, X : n j 

protokirki, V : 145 

protomanica, III : 91, X : 107, 115 

protonothodes, VII : 203 

proximans, I : 234 

pseudocarpenteri, V : 124 

pseudophaeus, III : 89, X : 107, 115 

pseudosmaragdalis, V : 89, VIII : 256 

publius, X : 75, 118 

purpurea, V : 141 

pnrpurina, VIII : 240 

pythodoris (us), I : 222, V : 79, X : 105, 113 

rabaiensis, VI : 202 

radiata, X : 81 

ragazzi, VI : 218 

rectans, VI : 229 

rectifascia, X : 76 

reducta Rothschild, II : 189 

reducta van Someren, VII : 214, X : 108 

regalis, V : 101 

regius, VIII : 228 

relalus, IX : 448 

richelmanni, VI : 236, X : no, 113 

rogersi, V : 131 

rosae, IV : 281, 284 

rosella, V : 140 

rosemariae, IX : 435 

ruandana, IV : 305 

rubiginea, X : 85 

rydoni, IV : 309, X : 104 



132 



V. G. L. van SOMEREN 



sabulosus, V : 156 

Sagittarius, X : 69 

saperanus, I : 210 

saturnalis, I : 203 

saturnus, I : 204 

schatzi, X : 87 

schoutedeni, II : 220, V : 79, X : 113 

schultzi, IX : 440 

selousi, VII : 222 

senegala, X : 103 

septentrionalis, I : 236 

seriata, V : 107 

shimbanus, I : 215 

smaragdalis, II : 210, X : 113 

somalicus, VI : 218 

splendens, V : 128 

stevensoni, I : 233 

stonehami, I : 218 

Stonehamia, X : 111 

subalbescens, IV : 312 

subargentea, V : 119 

subcaerulea, VIII : 240 

subornatus, IX : 440, X : 114 

subpallida, IX : 426 

suk, V : 132 

suna, X : 108 

superbus, IX : 436, X : no, 114 

suppressa, IV : 288 

swynnertoni, IV : 283 

talagagae, VIII : 218 

tanganika, VI : 199 

tanganyikae, III : 67, V : 80 

tavetensis, III : 62 

tectonis, I : 230, V : 78 

teita, I : 238 

teitensis, I : 216 

tenuis, VII : 219 

theresae, VI : 242 

thomasius, IX : 470 

thyestes, II : 189 

thysi (i), VIII : 219, X : 109, 115 

tiberiella, X : 90 

tiberius, X : 90, 118 

tiridates, VIII : 235, X : 113 

tiridatinus, VIII : 237 

toro, II : 214 



toyoshimai, IX : 459 
trajanus, X : 87, 118 
tricolor, V : 140 
tristis, VIII : 237 

ugandensis, VIII : 248 
ugandicus, VIII : 225 
ungemachi, VIII : 235, IX : 483 
usambarae, III : 73, X : 105, 115 
usambarensis, I : 214 
ussheri, X : 70, 118 

vansomereni, (Charaxes), V : 129 
vansomereni, (Euxanthe), X : 88 
vansoni, V : 137, X : 108, 117 
vansonoides, V : 145 
varanes, IX : 476, X : in 
variata, V : 138, X : 108, 117 
velox, IX : 470 
vetula, V : 88 
victoriae, VIII : 231 
viola, V : 127, X : 108, 116 
violetta, III : 50, X : 113 
violinitens, X : 73, 118 
violinitincta, V : 145 
viridicaerulea, V : 141 
viridicostatus, IX : 467 
virilis, V : 92, X : 115 
vologeses, IX : 478 
vumba, I : 211 
vumbui, II : 194 

wakefieldi, X : 85, 117 
wernickei, II : 203 
whitei, X : 105 
whytei, VII : 222 
woodi, II : 195, VIII : 259 

xiphares, II : 188, V : 82, VIII : 259, 
X :ii 3 

zambiae, VI : 246 
zelica, IX : 457, X : 117 
zingense, IV : 313 
Zingha, X 

zingha, IX : 466, X : 114 
zoippus, IX : 474 
zoolina, IX : 444, X : 114 



v. g. l. van someren 

The Sanctuary 

Ngong 

P. O. Box 24947 

Karen 

Kenya 



REVISIONAL NOTES ON AFRICAN CHARAXES 



133 




Map 1. 



134 



V. G. L. van SOMEREN 




dcctus 

violinitcns B 
violinitens bnamba □ 

ussheri usshen £ 

ussheri intcrposita $ 

iwsJieri <fofc«//i 

publius publius A 

publius (cntralis X 
publhif kigoma "^t- 



, "XX"" I 



Map 2. 



REVISIONAL NOTES ON AFRICAN CHARAXES 



135 




curinomt eurtnome 
eunnome ansclllCtt 
tunnome teledon 
tunnome tlgonue 
tunnome bltbirict 

CTMileyl cnssltyl 
'ssleyl an\org*i 
aossU/i magnlfi 
cnaleyl claudi 
madagaitaritri 
wakefltldl 



Map 3. 



PLATE I 

Palla 

Upper and undersides 

Figs i, 2. decius (Cramer), <$ (Sierra Leone: Port Lokko). Photos BMNH. 
Figs 3, 4. decius (Cramer), $ (Sierra Leone). Photos BMNH. 

Figs 5, 6. ussheri ussheri (Butler), $ Type (Gold Coast [Ghana]). Photos BMNH No 
50108-9. 

Fig. 7. ussheri ussheri (Butler), £ (Gold Coast [Ghana]). 
Fig. 8. ussheri ussheri (Butler), $ (Ivory Coast). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 1 










PLATE 2 



Palla ussheri (Butler) 

Upper & undersides 
Fig. g. ussheri g (Togo). 
Fig. 10. ussheri § (Togo). 

Fig. 11. ussheri £ form transitional to interposita Joicey & Talbot (East Congo: Epulu). 
Fig. 12. ussheri $ form transitional to interposita Joicey & Talbot (East Congo: Epulu). 
Fig. 13. ussheri 9 intermediate form (East Congo : Epulu). 
Fig. 14. ussheri $ dark var. (Moyen Congo [Congo: Brazzaville]). 
Fig. 15. ussheri £ intermediate form (Moyen Congo [Congo: Brazzaville]). 
Fig. 16. ussheri $ transitional to interposita Joicey & Talbot, resembling £ interposita of 
Uganda (East Congo : Beni). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 2 





lb 



O 





12 




hi 





14 






16 



PLATE 3 

Palla ussheri (Butler) 
Upper & undersides 

Fig. 17. ussheri <£ transitional to interposita retaining facies of the more western $ (East 
Congo: Beni). 

Fig. 18. interposita Joicey & Talbot, <$ (East Uganda). 

Figs 19, 20. ussheri § form dobelli Hall, Type (Cameroons [Cameroun]: Bitje, Ja River). 
Photos BMNH Nos 501 14-5. 

Figs 21, 22. interposita Joicey & Talbot, q* Type (Uganda: Mabira Forest). Photos BMNH 
Nos 50110-1. 

Figs 23, 24. interposita Joicey & Talbot, § (Uganda: Jinja). Photos BMNH Nos 50112-3. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 3 






11'// K . M 





20 







PLATE 4 

Palla 
Upper & undersides 

Fig. 25. ussheri interposita Joicey & Talbot, § (East Uganda). 

Fig. 26. violinitens violinitens (Crowley), g (Ivory Coast). 

Fig. 27. violinitens violinitens (Crowley), 9 (Ivory Coast). 

Fig. 28. violinitens cline to coniger (Butler), £ (Congo: Kasai). 

Fig. 29. violinitens cline to coniger (Butler), 9 (Congo: Kasai). 

Fig. 30. violinitens cline to coniger (Butler), Q (Central African Republic : Bangui) (/. 
Plantrou) . 

Figs 31, 32. violinitens cline to coniger (Butler), $ (Moyen Congo [Congo: Brazzaville]) 
(Jackson coll.). 



Bull. Br. Mus. nal. Hist. (Ent.) 32, 3 



PLATE 4 







28 






PLATE 5 

Palla 
Upper & undersides 

Fig. 33. violinitens cline to coniger (Butler), § (Moyen Congo [Congo: Brazzaville]). 

Fig. 34. violinitens £ transitional to bwamba subsp. n. (Eastern Congo: Epulu). 

Fig. 35. violinitens bwamba subsp. n., <$ holotype (W. Uganda: Bwamba Valley, v. 1954) 
{van Someren) . 

Fig. 36. violinitens bwamba subsp. n., $ paratype (W. Uganda: Bwamba Valley). 

Fig. 37. violinitens bwamba subsp. n., $ allotype (W. Uganda: Bwamba Valley, vii. 1942) 
(van Someren) . 

Fig. 38. publius publius Staudinger, $ (Nigeria: Old Calabar). 

Fig. 39. publius publius Staudinger, 9 (Ghana: Juaso). 

Fig. 40. publius centralis subsp. n., $ paratype (Moyen Congo [Congo: Brazzaville]) (Jackson 
coll.). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 5 










f 




40 




PLATE 6 

Palla publius Staudinger 
Upper & undersides 

Figs 41, 42. centralis subsp. n., <§ holotype (Afriq. Occid: Station Kamerun). 

Figs 43, 44. centralis subsp. n., 9 allotype (Cameroun). 

Fig. 45. centralis subsp. n., <$ paratype (Moyen Congo [Congo: Brazzaville]) (Jackson 
coll.). 

Fig. 46. kigoma subsp. n., g holotype (Tanzania: Kigoma, Kabogo, 28.xi.1961) (Japanese 
Primate Exped.). 

Figs 47, 48. kigoma subsp. n., g paratype (Tanzania: Kigoma, Kabogo, 28.xi.1961) (Japanese 
Primate Exped.). 

Figs 49, 50. publius f. moderata Gaede, g (Congo: Kapulumbo, Kasai). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 6 




PLATE 7 

Euxanthe eurinome (Cramer) 

Upper & undersides 

Fig. 51. eurinome, £ (Sierra Leone). 

Fig. 52. eurinome, 9 (Sierra Leone). 

F IG - 53- 54- eurinome, $ (Ivory Coast). 

Figs 55, 56. ansellica (Butler), <$ type (Angola: Kinsembo). Photos BMNH Nos 51244-5. 

Fig. 57. ansellica var. radiata van Someren & Rogers (C. Uganda: Mawakota). 

Fig. 58. ansellica (Butler), $ (Uganda: Bwamba Valley). 



Bull. Br. Mus. nal. Hist. (Ent.) 32, 3 



PLATE 7 




PLATE 8 

Euxanthc eurinome (Cramer) 
Upper & undersides 

Fig. 59. ansellica (Butler), £ (Uganda: Toro, Mpanga Forest). 

Fig. 60. ansellica (Butler), $ (Southern Cameroun). 

Fig. 61. ansellica (Butler), $ (Uganda: Bwamba Valley). 

Fig. 62. ansellica (Butler), $> (Uganda: Toro, Mpanga Forest). 

Fig. 63. ansellica (Butler), <j (Uganda: Bwamba Valley). 

Fig. 64. ansellica f. burgeoni Le Cerf, <$ holotype (E. Congo: Kindu). Photos MNHN, Paris. 
No underside figured. 

Fig. 65. celadon Le Cerf, $ holotype (Gabun). Photo MNHN, Paris. No underside 
figured. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 8 




PLATE 9 

Euxanthe eurinome (Cramer) 

Upper & undersides 

Figs 66, 67. elgonae subsp. n. $ paratypes (Kenya: Mara District, Gori River Forest). 
Fig. 68. elgonae subsp. n. $ paratype (Kenya: S.E. Mt Elgon, Trans Nzoia). 
Fig. 69. elgonae subsp. n. <§ paratype (Kenya: S.E. Mt Elgon, Trans Nzoia). 
Fig. 70, 72. elgonae subsp. n. £ holotype (Kenya: S.E. Mt Elgon, Trans Nzoia). 
Fig. 71, 73. elgonae subsp. n., ^ allotype (Kenya: S.E. Mt Elgon, Trans Nzoia). 



Bull. Br. Mus. nal. Hist. (Ent.) 32, 3 



PLATE 9 




PLATE 10 

Euxanthe crossleyi (Ward) 

Upper & undersides 

Fig. 74. crossleyi <J (E. Congo: Beni). 

Fig. 75. crossleyi $ (Moyen Congo [Congo: Brazzaville]). 

Figs 76, 77. crossleyi g type (Cameroun) . Photos BMNH Nos 501 16-7. 

Figs 78, 79. crossleyi $ type (Cameroun). Photos BMNH Nos 501 18-9. 

Fig. 80. crossleyi $ (Moyen Congo [Congo: Brazzaville]). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 10 




PLATE ii 

Euxanthe crossleyi (Ward) 
Upper & undersides 

Figs 8i, 83. ansorgei Rothschild & Jordan, $ (Uganda: Masaka District, west side of Lake 
Victoria north of Kagera River, Katera Forest). 

Fig. 82. ansorgei Rothschild & Jordan, $ (Uganda: Masaka District, west side of Lake 
Victoria north of Kagera River, Katera Forest). Note abnormal venation of hind wing. 

Fig. 84. ansorgei Rothschild & Jordan, <$ (Tanzania: Kigoma District). 

Fig. 85. ansorgei Rothschild & Jordan, g (Uganda: Kigezi District, Kayonza, lower 
Impenetrable Forest). 

Fig. 86. ansorgei Rothschild & Jordan, g (Kenya: Nandi-Elgon Forests, Kakamega). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 11 




PLATE 12 

Euxanthe crossleyi (Ward) 
Upper & undersides 

Fig. 87. ansorgei Rothschild & Jordan, <J (Uganda: Kigezi District, Kayonza, lower 
Impenetrable Forest). 

Fig. 88. ansorgei Rothschild & Jordan, § (Kenya: Nandi-Elgon Forests, Kakamega). 

Fig. 89. ansorgei § transitional to magnifica Rebel (Uganda: Masaka District, Katera 
Forest) . 

Figs 90, 91. magnifica Rebel, £ holotype (E. Congo: Nawambi-Irumu). Photos BMNH 
Nos 51246-7. 

Fig. 92. magnifica Rebel, g (Congo: Middle Lowa Valley, Walikali). 

Fig. 93. ansorgei Q transitional to magnifica Rebel (Uganda: Masaka District, Katera 
Forest) . 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 12 




PLATE 13 

Euxanthe 

Upper & undersides 

Figs 94, 95. wakefieldi (Ward), $ type (E. Africa: Ribe). Photos BMNH Nos 51256-7. 
Figs 96, 97. wakefieldi (Ward), <j> (Kenya: coastal forests). 
Figs 98-100. wakefieldi (Ward), $ (Kenya: coastal forests). 
Fig. ioi. madagascariensis (Lucas), <$ (Madagascar). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 13 





PLATE 14 

Enxanthe 
Upper & undersides 

Fig. 102. madagascariensis (Lucas), § (Madagascar). 

Fig. 103. trajanus trajanus (Ward), <j (Kamerun [Cameroun]). 

Fig. 104. tyajanus trajanus (Ward), § (Kamerun [Cameroun]). 

Fig. 105. trajanus cline to vansomereni Poulton, $ (Eastern Congo: Beni-Irumu). 

Fig. 106. trajanus vansomereni Poulton, g paratype (Uganda: Entebbe). Photos BMNH 
Nos 51252-3. 

Fig. 107. trajanus vansomereni Poulton, $ paratype (Uganda: Entebbe). Photos BMNH 
Nos 51254-5. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 14 




106 



107 



PLATE i 5 

Euxanthe trajanus (Ward) 

Upper & undersides 

Fig. 108. vansomereni Poulton, g (Uganda: Mawakota). 

Fig. 109. vansomereni Poulton, 9 (Uganda: Mawakota). 

Fig. no. gabonicus Le Cerf, $ type (Gabon, Echibanga). Photos BMNH Nos 51250-1. 

Fig. hi. gabonicus Le Cerf, g (Gabun). 

Fig. 112. antonius Rousseau-Decelle, <J paratypc (Congo: Kafakumba, Katanga). Photos 
I. Grahame. 

Figs 113, 114. nigenae subsp. n., g holotype (Nigeria: Ikom, Ogolo Prov.) {Jackson). 



Bull. By. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 15 




PLATE 16 

Euxanthe 
Upper & undersides 

Figs 115, 116. trajanus nigeriae subsp. n., § allotype (Nigeria: Ikom, Ogolo Prov.) (Jackson). 

Figs 117, 118. tibeyius tiberius Grose-Smith, $ holotype (Kenya: Mombasa). Photos 
BMNH Nos 5 1 260-1. 

Figs 119, 120. tiberius tiberius Grose-Smith, 9 allotype (Kenya: Mombasa). Photos 
BMNH Nos 51262-3. 

Fig. 121. tiberius tiberius Grose-Smith, <J (Kenya: coastal forests). 

Fig. 122. tiberius tiberius Grose-Smith, $ (Kenya: coastal forests). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 16 




PLATE 17 



Euxantlie tiberius Grose-Smith 

Upper & undersides 

Figs 123, 124. meruensis van Someren, g type (Kenya : Meru). Photos BMNH Nos 51268-9. 
Figs 125, 126. meruensis van Someren, $ type (Kenya: Meru). Photos BMNH Nos 51270-1. 
Fig. 127. meruensis van Someren, <-J (Kenya: Mt Kenya, Meru Forest). 
Fig. 128. meruensis van Someren, § (Kenya: Mt Kenya, Meru Forest). 

Figs 129-136. tiberius. Early stages from ova laid 25.vii.1964, obtained from a fy collected 
in the Makadara Forest. 

Fig. 129. First instar larva, 3 # ix.i964. 
Fig. 130. Second instar larva, 6.ix.ig64. 
Fig. 131. Third instar larva, 11.ix.1964. 
Fig. 132. Fourth instar larva, 15. ix. 1964. 
Fig. 133. Fifth instar larva, 26.ix.1964. 
Fig. 134. Larval heads from the five instars. 
Figs 135, 136. Pupae, i.x.1964. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 17 




PLATE iS 

Charaxes 
Upper & undersides 

Figs 137, 138. octavus Minig = patergodarti Neidhoefer, $ holotype (Central African 
Republic: Bangui). Photos J. Neidhoefer. 

Figs 139, 140. ansorgei rydoni van Someren, $ neallotype (Tanzania: Usambara Mts, 
Lushoto, Magamba, 6000 ft, 4.^.1973) (/. Bampton). (From coloured photos by W. H. 
Henning.) 

Figs 141, 142. eudoxus lucyae subsp. n., $ holotype (Tanzania: Usambara Mts, Magamba 
Forest, Lushoto, 6000 ft, 4.^.1973) (I. Bampton). (From coloured photos by W. H. Henning.) 

Figs 143, 144. eudoxus lucyae subsp. n., 9 paratype (Tanzania: Magamba Forest, Lushoto, 
5000 ft, ii. 1974) ( s - c - Collins). Photos BMNH. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE ii 





PLATE 19 

Charaxes 
Upper & undersides 

Figs 145, 146. cithaeron cithaeron $ aberration whitei ab. n., holotype (South Africa: Natal, 
Eshowe, 8.V.1971) (R. S. White). Photos BMNH. 

Figs 147, 148. usambarae $ form collinsi forma n., holotype (Tanzania: Usambara Mts, 
Amani, x. 1973) (S. C. Collins). Photos BMNH. 

Figs 149, 150. martini van Someren, $ (Malawi: Mlange Mt, Malosa Stream, 15.x. 1971) 
(C. H. McCleery). 

Figs 151, 152. ma/w^a van Someren, $ neallotype (Uganda: Ruhiza, Impenetrable Forest, 
8000 ft, 28. ix. 1972) (I. Bampton). Photos BMNH. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 3 



PLATE 19 










ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172 : 500 text -figures. September, 

1965- £3-25- 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3-i5. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae) . Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 
68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. 
February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. 

21." 'Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text-figures. December, 1973. £9.55. 

printbd by Unwin Brothers Limited thb grbsham press old woking surrey England 






THE SPECIES OF CHALCIDOIDEA 
DESCRIBED FROM NORTH AMERICA 

NORTH OF MEXICO BY 
FRANCIS WALKER (HYMENOPTERA) 



B. D. BURKS 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 4 

LONDON : 1975 



THE SPECIES OF CHALCIDOIDEA DESCRIBED 

FROM NORTH AMERICA NORTH OF MEXICO 

BY FRANCIS WALKER (HYMENOPTERA) 



BY 

BARNARD De WITT BURKS 

Systematic Entomology Laboratory, Agricultural Research 
Service, U.S.D.A., Washington, D.C. 20250, U.S.A. 



Pp. 137-T-7 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 4 

LONDON : 1975 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Departments 
of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 4 of the Entomological 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Ent.) 



ISSN 0524-6431 



Trustees of the British Museum (Natural History), 1975 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 28 July, 1975 Price £2.45 



THE SPECIES OF CHALCIDOIDEA DESCRIBED 

FROM NORTH AMERICA NORTH OF MEXICO 

BY FRANCIS WALKER (HYMENOPTERA) 

By B. D. BURKS 



CONTENTS 



Synopsis 

Introduction 

Acknowledgements 

Eulophidae 

Encyrtidae . 

eupelmidae . 

Perilampidae 

torymidae 

Ormyridae 

Pteromalidae 

Eurytomidae 

Chalcididae 

Leucospididae 

References . 

Index 



Page 

J 39 
*39 

141 
141 

147 
149 

150 
151 
153 
154 
162 
164 
166 
166 
169 



SYNOPSIS 

The 72 species of Chalcidoidea described by Francis Walker from North America north of 
Mexico are reviewed and full information is given on their type-material. The species are 
assigned, where possible, to their currently correct genera, and synonymies are given. Twenty- 
six specific names are newly synonymized. Sixty-three lectotypes are newly designated, of 
which eight are for other authors' species here synonymized with those of Walker. 



INTRODUCTION 

Francis Walker described 72 species of Chalcidoidea from North America north 
of Mexico. Of these, 59 were described from collections made by E. Doubleday 
at St John's Bluff, Florida. Another species, described from 'East Florida, Double- 
day,' may also have been from St John's Bluff, and another, from 'North America, 
Doubleday' might have been from St John's Bluff. Five other species were des- 
cribed from specimens collected by G. Barnston at St Martin's Falls, Albany River, 
Ontario, Canada. Two species were from collections John Abbot made in Georgia. 
There was one species each from the collections of W. S. M. D 'Urban in Canada 
and R. Forster in Ohio. One species was described from a specimen reared in 



i 4 o B. D. BURKS 

England from a pine cone from California, and another was from a specimen taken 
by an unknown collector 'near New York.' 

St John's Bluff, where Doubleday collected the bulk of these Walker species, 
is a locality still preserved, in Duval County, Florida. It is located on the south 
bank of the St John's River and is now contained in the Fort Caroline National 
Memorial. This is approximately 12 kilometres east of the city of Jacksonville, 
and about 8 kilometres inland from the Atlantic Ocean. It lies between 8i° 29' 
and 8i° 30' W at 30 23' 15" N. The bluff itself rises 12-5 metres from the river 
bank, just east of the remains of Fort Caroline. It is approximately 77-5 metres 
from the river bank to the crest of this bluff, and the bluff extends approximately 
930 metres from east to west. The bluff and the immediately adjacent land has 
a vegetative cover. Approximately 310 metres south of the crest of St John's 
Bluff the land becomes marshy. 

Barnston collected his specimens at St Martin's Falls on the Albany River, 
in Ontario, Canada. This locality is often cited as Hudson Bay, but it is actually 
far inland from Hudson Bay. St Martin's Falls, which was at one time a Hudson 
Bay Company post, is on the Albany River some 80 kilometres upstream from the 
confluence with the Ogoki River. This is at about 51 ° 30' N, 86° 30' W, approxi- 
mately 440 kilometres upstream from Fort Albany on James Bay. 

The balance of the species Walker described from North America north of Mexico 
are from scattered localities about which no further data are available. 

In this paper each Walker species is assigned, as far as possible, to its currently 
correct genus, and synonymies are given. In the few cases where a Walker name is 
synonymized with an earlier name (either of Walker or of another author) it is 
dealt with under the valid name. The species are listed separately for each family 
and, within each family, they are listed in alphabetical order of the valid specific 
name. 

Of the 72 Walker species, lectotypes for 4 species have already been designated, 
and the types of 4 others are missing. The types of all the rest are in the British 
Museum (Natural History) (hereafter abbreviated to BMNH). In this paper I 
have designated lectotypes for 63 species. The circular colour-coded BMNH type- 
labels have been added to all the Walker types examined, purple for lectotypes 
and blue for paralectotypes. Each lectotype bears in addition a label stating 
'LECTOTYPE [name of species] designated B. D. Burks 1975'. I refrained from 
designating a lectotype for one species because it is now represented in the collection 
by a single, very poor specimen. There is a chance that a better type-specimen 
of this species will be found eventually. I give here all the data that are on the 
Walker types. Some specimens have no label data, others carry only the Walker 
species name. 

In this paper I have also designated lectotypes for several species described by 
other authors, the types of which are in the U.S. National Museum (hereafter 
abbreviated to USNM). This was required because those species, represented 
by syntypes, are here synonymized under Walker species. 

In general the Walker types of North American Chalcidoidea are quite well 
preserved, although all of them are over a century old, and most of them are about 



SPECIES OF CHALCIDOIDEA 141 

140 years old. There has not been so much colour change in them as I would have 
expected. Their colours are not greatly different from the colours of specimens 
of the same species that I have seen that were collected recently. 

These Walker types were probably all originally pinned into the BMNH collection, 
either mounted on cards and with the wings spread, or mounted on small pins 
that were set on small cards. About 1900 the entire collection was repinned and 
relabelled by C. 0. Waterhouse. He added name labels to the type-specimens; 
on the reverse side of each label there is the statement 'Stood under this name in 
old B. M. Coll. V. Waterhouse.' There is every indication that this relabelling was 
done with the greatest care, but in many species the Waterhouse label does not give 
the name exactly as it was in the original description. This may be due to the 
fact that the labels were changed in the old collection during the approximately 
55 years the specimens were there, or the name labels in the collection may not 
have agreed with the published names from the beginning. At any rate it would 
have been better to have preserved all of Walker's original name labels, but this 
was done for only a few species. 

About 1930 nearly all the Walker types of Chalcidoidea were remounted onto 
card points by G. B. Thompson. The results were unfortunate for many species. 
Several of these types have the legs and wings entangled in the mounting medium. 
It is difficult to make out the characters of the appendages in these types. In 
others an excessive amount of the mountant was used. However, it has been 
possible to remount some of these successfully. 



ACKNOWLEDGEMENTS 

I am greatly indebted to Dr Z. Boucek, Commonwealth Institute of Entomology, 
for giving me his opinions on several of these Walker type-specimens, the condition 
of which made placement difficult. Dr H. V. Weems, Florida Department of 
Agriculture and Consumer Services, Gainesville, kindly furnished information 
about St John's Bluff. Dr O. Peck, formerly of the Canada Department of Agri- 
culture, Ottawa, provided information about St Martin's Falls. 



EULOPHIDAE 

Notanisomorpha calavius (Walker) comb. n. 

Eulophus Calavius Walker, 1847 : 24, 9> 6*- 
Hemiptarsenus calavius (Walker) Peck, 195 1 : 428. 

Type-material. There are two female and one male syntypes of this species 
in the BMNH collection on card points. The first specimen in the series bears 
the label 'Calavius' in Walker's hand. I designate the second female specimen, 
labelled 'B.M. Type Hym. 5. 2268', as LECTOTYPE. Described from St John's 
Bluff, Florida. 



i 4 2 B. D. BURKS 

Remarks. Although this is a very distinctly marked species, it does not seem 
to have been redescribed since Walker's time. There is a series of specimens from 
Maryland in the USNM collection. These specimens had long been thought to 
represent an undescribed species. 

Distribution. Maryland, Florida. 

Necremnus cyriades (Walker) 

Eulophus Cyriades Walker, 1847 : 25, £. 
Necremnus cyriades (Walker) Peck, 195 1 : 428. 

Type-material. One male specimen on a card point is in the BMNH collection; 
it is in poor condition. It is labelled 'B.M. Type Hym. 5. 2269' and 'Cyriades' 
in Walker's hand. I designate it LECTOTYPE. The right antenna is broken 
off beyond the pedicel, but the left antenna still retains funiculars 1 to 3 which 
bear long branches. Described from St John's Bluff, Florida. 

Remarks. There is a single male specimen of this species from Washington, 
D.C. in the USNM collection. This is the one mentioned by Chittenden (1901 : 86). 

Distribution. District of Columbia, Florida. 

Biology. Has been reared from the oleuthreutid moth, Hedya ochroleucana 
(Frolich) . 

Achrysocharella damastes (Walker) comb. n. 

Closterocerus Damastes Walker, 1847 : 22, 9- 

Entedon Damastes (Walker) Walker, 1848 : 137; Peck, 1951 : 470. 

Type-material. One female specimen, mounted on a card, is in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 2270' and 'Entedon Damastes Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. 

Remarks. This is a brilliantly coloured and distinctively marked species, yet 
it does not seem to have been redescribed since Walker's time. 

Distribution. Florida. 

Aprostocetus epidius (Walker) comb. n. 

Tetrastichus Epidius Walker, 1847 : 28, $. 

Aprostocetus granulatus Ashmead, 1888 : 105, ?J- Syn. n. 

Type-material. Tetrastichus epidius Walker. There are two female speci- 
mens on card points in the BMNH Collection, one of which is labelled 'B. M. Type 
Hym. 5. 1413; Type, C. F. 1938' and 'Epidius' in Walker's hand. I designate 
this specimen LECTOTYPE. Described from St John's Bluff, Florida. 

Aprostocetus granulatus Ashmead. Described from one female specimen from 
Jacksonville, Florida; the holotype is USNM no. 28662. 



SPECIES OF CHALCIDOIDEA 143 

Remarks. I left this species unplaced in my revision of the North American 
species of Tetrastichus (Burks, 1943 : 604), on the basis of the notes that Dr Ferriere 
had sent me about the type. When I saw the type myself, however, I recognized 
it as the species that has been called Aprostocetus grantdattts Ashmead in the North 
American literature for almost 80 years. 

Distribution. Ontario south to Florida, west to Illinois. 

Biology. Reared from dipterous gall on Sambucus. 

Cirrospilus eunapius Walker 

Cirrospilus Eunapius Walker, 1847 : 27, 9- 

Type-material. Three female syntypes on card points are in the BMNH 
collection, the first in the series labelled 'Eunapius' in Walker's hand. I designate 
the second specimen, labelled 'B.M. Type Hym. 5 2271', as LECTOTYPE. 
Described from St John's Bluff, Florida. 

Remarks. This is a distinctive species, but I have not seen any specimen of 
it except these types. These are a brownish purple (may be somewhat faded), 
with a broad yellow band crossing the praescutum and all but the posterior tips 
of the two segments of the mesoscutum. There is another yellow cross band at 
the base of the gaster. The forewing has a broad brown band extending from the 
marginal vein to the posterior wing margin. The apices of the coxae, trochanters, 
and the bases of the femora are white, the legs otherwise dark brown. The antenna 
has the first funicular segment slender, one and one-third times as long as the broader 
second funicular; the club is as long as the funiculus. The scutellum is quite flat 
and smooth, with the sublateral carinae distinct. The propodeum has a strong 
longitudinal, median carina. This well-marked species should be easily identified. 

Distribution. Florida. 

Tetrastichus gala Walker 

Tetrastichus Gala Walker, 1847 : 28, 9. 

Type-material. One specimen on a card is in the BMNH collection. It is 
labelled 'B. M. Type Hym. 5. 1412; Type, C. F., 1938' and 'Tetrastichus Gala 
Walker' by Waterhouse. I designate it LECTOTYPE. Described from St John's 
Bluff, Florida. 

Remarks. There are specimens of this species, all from Florida, in the USNM 
collection. 

Distribution. Florida. 

Tetrastichus granulatus Walker 

Tetrastichus granulatus Walker, 1844 : 17, $. 

Type-material. There are three female syntypes and the gaster of a fourth 
female in the BMNH collection. One specimen, mounted on a card, is labelled 



144 B. D. BURKS 

'B. M. Type Hym. 5. 1411; 44.17, Hudson's Bay, St. Martin's Falls' and 'Tetrastichus 
granulatus Walker' by Waterhouse, also 'Type C. F. 1938.' I designate it LECTO- 
TYPE. One of the other specimens is a different species. 

Remarks. I have not seen this species of Tetrastichus before, but it should be 
possible to identify it. The head and body are metallic greenish black, the femora 
dark metallic green, the tibiae yellow, the antennae dark brown. The forewing 
has 2 submarginal bristles and the marginal vein is only twice as long as the stigmal 
vein. Each lateral margin of the praescutum has a single sparse row of bristles. 
The propodeum is smooth and ecarinate. The ovipositor sheaths do not extend 
to the apex of the gaster. 

Distribution. Ontario. 



Galeopsomyia haemon (Walker) 

Tetrastichus Haemon Walker, 1847 : 28, ^. 
Horismenus haemon (Walker) Burks, 1943 : 605. 
Galeopsomyia haemon (Walker) Burks, 1971 : 82. 
Euderus columbiana Ashmead, 1888 : 104, 9. Syn. n. 
Galeopsomyia columbiana (Ashmead) Girault, 19 16 : 348. 

Type-material. Tetrastichus haemon Walker. There are two female and two 
male syntype specimens; one female is labelled 'B. M. Type Hym. 5. 1414; Type 
C. F., 1938' and 'Haemon' in Walker's hand. I designated it lectotype in an earlier 
paper (Burks, 1971 : 82). Described from St John's Bluff, Florida. This species 
is the same as Euderus columbiana Ashmead, the one Girault made type-species 
of his genus Galeopsomyia. 

Euderus columbiana Ashmead. This was described from an unspecified number 
of female specimens from the District of Columbia and Florida. There are now 
six syntype specimens under USNM no. 19923. One, labelled '3192°, Aug. 13, 
83; Euderus columbiana Ashm.; Galeopsomyia genotype' is designated LECTO- 
TYPE. USNM records show that 3192 was the District of Columbia. 

Distribution. New York south to Florida west to Missouri and Arizona. 

Biology. Emerges from cecidomyiid galls. 



Euderus herillus (Walker) 

Entedon (Euderus) Herillus Walker, 1847 : 23, 9- 

Euderus herillus (Walker) Dalla Torre, 1898 : 6. 

Euderus (Leipocrossus) herillus (Walker) Yoshimoto, 1971 : 556. 

Type-material. Two female specimens, one of which, designated lectotype by 
Yoshimoto (1971 : 558), is labelled 'B. M. Type Hym. 5. 2091.' Described from 
St John's Bluff, Florida. 

Distribution. Virginia, Georgia, Florida. 



SPECIES OF CHALCIDOIDEA 145 

Neochrysocharis imbrasus (Walker) comb. n. 

Entedon Imbrasus Walker, 1847 : 23, $. 
Encyrtus imbrasus (Walker) Dalla Torre, 1898 : 259. 
Microterys imbrasus (Walker) Ashmead, 1900 : 391. 
Chrysocharis imbrasus (Walker) Graham, 1963 : 234. 

Type-material. One female specimen mounted on a card is in the BMXH 
collection, labelled 'B. M. Type Hym. 5. 2274' and 'Entedon Imbrasus Walker' by 
Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, Florida. 

Distribution. Florida. 

Necremnus iphinoe (Walker) 

Eulophus Iphinoe Walker, 1847 : 25, g. 
Necremnus iphinoe (Walker) Peck, 1951 : 428. 
Eulophus Gobryas Walker, 1847 : 26, ',$' = £. Syn. n. 

Type-material. Eulophus iphinoe Walker. One male specimen on a card 
point is in the BMNH collection, labelled 'B. M. Type. Hym. 5. 2272' and 'Iphinoe' 
in Walker's hand. I designate it LECTOTYPE. Described from St John's 
Bluff, Florida. It clearly is the male that should be associated with the female 
Walker described under the name Eulophus gobryas. 

Eulophus gobryas Walker. One female specimen on a card point is in the BMNH 
collection, labelled 'B. M. Type Hym. 5. 2273' and 'Gobryas' in Walker's hand. I 
designate it LECTOTYPE. Described from St John's Bluff, Florida. 

Distribution. Florida. 

Stenomesius levana (Walker), comb. n. 
Elachestus Levana Walker, 1847 : 27, §. 

Type-material. There are two female specimens, mounted on card points, 
in the BMNH collection. The first of these, labelled 'B. M. Type Hym. 5. 2275' 
and 'Levana' in Walker's hand, is designated LECTOTYPE. 

Remarks. This species does not seem to have been collected again since the 
original specimens were taken on St John's Bluff. Its characters place it in the 
genus Stenomesius. It has a well marked occipital carina; the pronotum is promin- 
ently carinate anteriorly; the thorax is strongly sculptured dorsally; the scutellum 
is carinate laterally and posteriorly, this carina formed by a series of minute, elongate, 
pits; the propodeum has two median carinae, these converging to form a single one 
near centre of the propodeum; the gastral petiole is elongate, the gaster semi-ovate. 

Distribution. Florida. 

Pnigalio minio (Walker) 

Eulophus Minio Walker, 1847 : 26, $. 

Pnigalio minio (Walker) Peck, 1951 : 426. 

Elachistus proximus Ashmead, 1894 : 34°> ?• Syn. n. 



146 B. D. BURKS 

Type-material. Eulophus minio Walker. There are two female specimens 
on card points in the BMNH collection under this name. The first is a Pnigalio 
labelled 'B. M. Type Hym. 5. 2276' and 'Minio' in Walker's hand. I designate 
it LECTOTYPE. The second specimen belongs in another genus. Described 
from St John's Bluff, Florida. Miller (1970 : 78) left this species unplaced, in 
his revision of the North American species of Pnigalio. 

Elachistus proximus Ashmead. Described from a single female specimen; the 
holotype is USNM no. 23452. Described from Morgantown, West Virginia. Miller 
(1970 : 12) gives several synonyms under proximus. 

Distribution. Quebec south to Florida west to the Mississippi Valley. 

Biology. A primary parasite of leaf-miners. 

Aulogymnus minyas (Walker), comb. n. 

Eulophus Minyas Walker, 1847 : 26, §. 

Type-material. There are two syntype female specimens in the BMNH collec- 
tion mounted on card points. The first specimen is labelled 'Minyas' in Walker's 
hand. I designate the second specimen, labelled 'B. M. Type Hym. 5. 2277', as 
LECTOTYPE. Described from St John's Bluff, Florida. 

Remarks. This species does not seem to have been found again since the original 
description. It clearly belongs in Aulogymnus, and is somewhat similar to the 
European A . aceris Foerster. 

Distribution. Florida. 

Acrias nileus Walker 

Acvias Nileus Walker, 1847 : 29, 9- 

Type-material. One female specimen mounted on a card is in the BMNH 
collection, labelled 'B. M. Type Hym. 5. 2278' and 'Nileus' in Walker's hand. 1 
designate it LECTOTYPE. Described from St John's Bluff, Florida. 

Remarks. This distinctively marked species, type-species of the genus Acrias 
Walker, has remained essentially unrecognized since its description over 130 years 
ago. Ashmead is supposed to have seen this type before he prepared his Classifica- 
tion of the Chalcid Flies (1904). If so, the characters he gives for it, 'head large, 
much wider than the thorax; wings with one or two fuscous fasciae; abdomen 
conic-ovate, the second segment occupying half the whole surface', are extremely 
misleading. In fact, the head is not large, but is the same width as the thorax; 
the forewing has a distinct longitudinal, dark streak in the basal half and peripheral 
dark spots in the apical area; the gaster is narrow and elongate; and the second 
gastral segment occupies only one-sixth the length of the gaster. The genus Acrias, 
as represented by its type-species, is closely related to Euderus Haliday and, thus, 
has only very remote relationship to such genera as Paracrias Ashmead or Eupara- 
crias Brethes. 



SPECIES OF CHALCIDOIDEA 



*47 



There is a specimen of Acrias nileus in the USNM collection from Willow Grove, 
Pa., reared 2 June, 1932, from a host that is thought to be Agonopterix nebulosa 
Zeller. This specimen has been labelled as an unknown genus of Eulophidae for 
nearly 40 years. 

Distribution. Pennsylvania, Florida. 



Horismenus sardus (Walker) 

Entedon [Horismenus) Sardus Walker, 1847 : 23, Cj- 

Encyrtus sardus (Walker) Dalla Torre, 1898 : 263. 

Pseudomphale sardus (Walker) Girault, 1918 : 130. 

Horismenus sardus (Walker) Peck, 1951 : 468. 

Elachristus flavipes Ashmead, 1886 : 133, £. [Synonymized by Burks, 1971 : 81.] 

Type-material. Entedon sardus Walker. One female specimen, mounted 
on a card, is in the BMNH collection, labelled 'B. M. Type Hym. 5. 2074; Horis- 
menus sardus Wlk. Type G. J. Kerrich det. 1969' and 'Sardus' in Walker's hand. 
I designate it LECTOTYPE. Described from North America. 

Remarks. This species has been treated in my recent revision of the North 
American species of Horismenus Walker (Burks, 1971). 



ENCYRTIDAE 

Pseudencyrtus bolus (Walker) comb. n. 

Encyrtus bolus Walker, 1844 : 17, $. 

(?) Microterys bolus (Walker) Ashmead, 1900 : 393. 

Microterys bolus (Walker) Peck, 195 1 : 488. 

Type-material. Three female specimens on individual card points were on 
a single pin in the BMNH collection, labelled 'Type; Spalangia from swelled willow 
twig; B. M. Type 5. 1, 115' and 'Encyrtus Bolus Walker' by Waterhouse. The 
second specimen of the three was repinned and labelled LECTOTYPE, and is here 
designated as such; the other two were labelled 'paralectotype'. The lectotype 
has both antennae intact and all legs and wings remaining, but is somewhat entangled 
in the mounting medium. Described from St Martin's Falls, Albany River, Ontario. 

Distribution. Ontario, Idaho. 

Biology. There is a series of this species in the USNM collection that was 
reared at Krassel, Idaho, from dipterous galls on Salix. 

Chrysopophagus flaccus (Walker) comb. n. 

Encyrtus (Cerchysius) Flaccus Walker, 1847 : 21, $. 
Cerchysius flaccus (Walker) Dalla Torre, 1898 : 244. 
Saranotum americanum Perkins, 1906 : 260, Cj. Syn. n. 
Chrysopophagus americanus (Perkins) Peck, 195 1 : 499. 



148 B. D. BURKS 

Type-material. Encyrtus flaccus Walker. A single female specimen, mounted 
on a card point is under the genus Cerchysius, labelled 'B. M. Type Hym. 5. 2279' 
and 'Cerchysius Flaccus Walker' by Waterhouse. I designate it LECTOTYPE. 
Described from St John's Bluff, Florida. This specimen is poorly mounted and the 
all important dorsal thoracic bristles have been rubbed off, but it clearly is the 
species long known in North American literature as americanus Perkins. 

Saranotum americanum Perkins. This was described from one female specimen 
from Ohio; the type is in the Bishop Museum, Honolulu, and I have not seen it. 
There are, however, specimens in the USNM that P. H. Timberlake compared 
with this type. 

Distribution. Continental United States, Hawaii. 

Biology. This is a secondary parasite of leafhoppers, emerging from the cocoons 
of their dryinid primary parasites. 

Encyrtus gastron Walker 

Encyrtus gastron Walker, 1847 : 21, §. 

Type-material. There is a single specimen, mounted on a card point, in the 
BMNH collection. It is labelled 'B. M. Type Hym. 5. 2280' and 'Gastron, Walk. 
Fla.' in Walker's hand. I designate it LECTOTYPE. Described from St John's 
Bluff, Florida. This specimen is so poorly preserved that it probably never can 
be placed. Both antennae are missing, the left forewing is missing, and the hind- 
wings and legs are embedded in the mounting medium. 

Distribution. Florida. 

Encyrtus reate Walker 

Encyrtus Reate Walker, 1847 : 22, <J. 
Type-material. Lost. Described from St John's Bluff, Florida. 
Distribution. Florida. 

Psyllaephagus vectius (Walker) comb. n. 

Encyrtus Vectius Walker, 1847 : 21, £. 
Syrphophagus vectius (Walker) Ashmead, 1900 : 398. 
Encyrtus solus Howard, 1885 : 15, $. Syn. n. 
Psyllaephagus solus (Howard) Gahan & Waterston, 1926 : 375. 

Type-material. Encyrtus vectius Walker. A single badly broken male speci- 
men, mounted on a card point, is in the BMNH collection. It is labelled 'B. M. 
Type Hym. 5. 2281' and 'Encyrtus Vectius Walker' by Waterhouse. I designate 
it LECTOTYPE. Described from St John's Bluff, Florida. The specimen is 
fragmentary, lacking all legs and all apical segments of both antennae, but the 
characteristic, deep-striate sculpture of the scutellum and the dark shadow in the 



SPECIES OF CHALCIDOIDEA 149 

anteroapical angle of the forewing will place this as the species long known as 
Psyllaephagus solus (Howard). 

Encyrtus solus Howard. Described from one female specimen from Gainesville, 
Florida; the holotype is USNM no. 2637. 

Distribution. Florida. 

Biology. Has been reared from the gall of Trioza magnoliae (Ashmead). 

EUPELMIDAE 

Eupelmella epicaste (Walker) 

Eupelmus Epicaste Walker, 1847 : 20, £. 
Eupelmella epicaste (Walker) Peck, 195 1 : 512. 
Macronettra epicaste (Walker) Peck, 1963 : 493. 
Eupelminus meteori Gahan, 1913 : 437, $. Syn. n. 
Eupelmella meteori (Gahan) Peck, 195 1 : 512. 
Macronettra meteori (Gahan) Peck, 1963 : 494. 

Type-material. Eupelmus epicaste Walker. There is a single female specimen, 
mounted on a card point, in the BMNH collection under Eupelmus, labelled 'B. M. 
Type Hym. 5. 2282' and 'Eupelmus Epicaste Walker' by Waterhouse. I designate 
it LECTOTYPE. Described from St John's Bluff, Florida. This specimen is 
intact, but evidently at some time had been broken in two at the propodeum and 
then rejoined. This is the species now being called meteori Gahan. 

Eupelminus meteori Gahan. Described from five female specimens; one specimen, 
labelled 'Brownsville, Tex., Exp. 11; Webster no. 6446 b 7 ; Type No. 16355 
U. S. N. M.' is designated LECTOTYPE. 

Distribution. Florida, Kansas, Texas. 

Biology. This is secondary parasite, emerging from the cocoons of braconid 
primary parasites. 

Eupelmus fonteia Walker 

Eupelmus Fonteia Walker, 1847 : 19, $. 

Type-material. There are three female syntypes in the BMNH collection 
under this name. The specimens are on card points, and each is labelled 'Eupelmus 
fonteia Walker' by Waterhouse. I designate the first specimen in the series, 
labelled 'B. M. Type Hym. 5. 2283', as LECTOTYPE. Described from St John's 
Bluff, Florida. 

Remarks. This is Eupelmus as that genus is currently understood, but this 
species cannot be placed until the genus is revised for North America. Eupelmus 
fonteia is one of the species lacking an exserted ovipositor, the head and body are 
iridescent coppery black, and the antennal scape is black; the body pubescence 
is silvery, the femora and tibiae are yellow-brown with slight darkening at the 
apices of the femora and middles of the tibiae; and tarsi are white. 

Distribution. Florida. 



150 B. D. BURKS 

Eupelmus lamachus Walker 

Enpelmus lamachus Walker, 1847 : 20, §. 

Type-material. There is a single specimen, mounted on a card point, in the 
BMNH collection labelled 'B. M. Type Hym. 5. 2284' and 'Eupelmus Lamachus 
Walker' in Waterhouse's hand. I designate it LECTOTYPE. It is in poor condi- 
tion. Described from St John's Bluff, Florida. 

Remarks. This is Eupelmus as the genus is currently understood, but this 
species cannot be placed until Eupelmus is revised. This is one of the thickset, 
large species of Eupelmus. It has the head and body red-coppery black (a colour 
that seems to occur only in Florida species). The midlegs are entirely yellow, 
but the other legs are yellow only beyond the coxae; the antennae are black; the 
compound eyes are large, and the lateral ocelli almost but not quite touch the 
margins of the compound eyes; the hind tibiae are not flattened and there are no 
visible bristles on the scutellum. 

Distribution. Florida. 

PERILAMPIDAE 

[Perilampus alexinus Walker 
See under Chalcididae below.] 

Perilampus hyalinus Say 

Perilampus hyalinus Say, 1828 : 79. 

Perilampus Entellus Walker, 18436 : 103, <j>, g. Syn. n. 

Type-material. Perilampus hyalinus Say. Type lost, but the identity of 
this species, originally described from Pennsylvania, has long been accepted on the 
basis of the identifications used by Smith (1912 : 34-48), Crawford (1914 : 70) 
and Smulyan (1936 : 380), all of which agree. 

Perilampus entellus Walker. There is one female specimen, mounted on a small 
pin on a card, in the BMNH collection, labelled 'B. M. Type Hym. 5. 2285' and 
'Perilampus Entellus' in Walker's hand. I designate it LECTOTYPE. This 
species was originally described from several specimens of both sexes from Ohio. 
The single remaining specimen runs to hyalinus Say in Smulyan (1936 : 372), 
and is in agreement with the current concept of this species. 

Distribution. North America. 

Perilampus lepreos Walker 

Perilampus lepreos Walker, 1846 : 89. 
Euperilampus lepreos (Walker) Peck, 1951 : 516. 

Type-material. Missing. Described from Georgia. 
Distribution. Georgia. 



SPECIES OF CHALCIDOIDEA 151 

TORYMIDAE 

Torymus aea (Walker) 

Callimome Aea Walker, 18436 : 104, 9- 

Torymus aea (Walker) Cresson, 1887 : 237. 

Torymus aeus (Walker) Peck, 1951 : 521. 

Syntomaspis amelanchieris Cushman, 1918 : 82, $. <$■ Syn. n. 

Torymus amelanchieris (Cushman) Peck, 1951 : 521. 

Type-material. Callimome aea Walker. There is a single female specimen, 
mounted on a small pin on a card, in the BMNH collection, labelled 'B. M. Type 
Hym. 5. 44' and 'Callimome aea Walker' by Waterhouse. There is also a locality 
label, 'St. John's Bluff,' added in error by someone after Walker's time; this species 
was described from near New York. I designate this specimen LECTOTYPE. 
This is the species described many years later by Cushman as amelanchieris. 

Syntomaspis amelanchieris Cushman. Described from five females and one male 
from Pickens, West Virginia; the holotype female is USNM no. 20968. 

Distribution. North-eastern North America. 

Biology. Milliron (1949 : 393-395) has shown that the larva of this species 
develops in an Amelanchier seed first as an external parasite on the larva of Megas- 
tigmus amelanchieris Cushman and, after having consumed it, becomes phytopha- 
gous and completes its development by feeding on the seed itself. This type of 
parasitic-phytophagous development occurs elsewhere in the Chalcidoidea (Maly- 
shev, 1968 : 53). 

Megastigmus albifrons Walker 
Megastigmus albifrons Walker, 1869 : 314, <$. 

Type-material. One male specimen, mounted on a card, is in the BMNH 
collection, labelled 'albifrons' in Walker's hand and 'B. M. Type Hym. 5.67.' I 
designate it LECTOTYPE. Reared in England from a pine cone collected in 
California. This certainly is the species currently being identified as albifrons. 

Distribution. South-western United States. 

Biology. Develops in the seeds of Pinus ponderosa. 

Torymus cecidomyiae (Walker) 

Callimome Cecidomyiae Walker, 1844 : 15, $. 
Torymus cecidomyiae (Walker) Cresson, 1887 : 237. 
Callimome strobiloides Huber, 1927 : 40, 9- Syn. n. 
Torymus strobiloides (Huber) Peck, 195 1 : 526. 

Type-material. Callimome cecidomyiae Walker. Two female specimens in 
poor condition are in the BMNH collection. Each is labelled 'Hudson's Bay; 
Callimome Cecidomyae Walker' by Waterhouse. I designate the second specimen, 
labelled 'B. M. Type Hym. 5. 2286', as LECTOTYPE. This specimen was originally 



152 B. D. BURKS 

on a pin; but it has been remounted on a card point. The other specimen has the 
thorax with the attached left forewing mounted on a card, the other fragments 
cemented to the locality label beneath. This is recognizably the same as Huber's 
species strobiloides. 

Callimome strobiloides Huber. Described from three female specimens from 
Columbus, Ohio. The holotype female is USNM no. 25351. 

Distribution. Quebec, Ontario, New Hampshire, New York, Ohio, Minnesota. 

Biology. Reared from midge galls on buds of Salix. 

Torymus lissus (Walker) 

Callimome lissus Walker, 1843a : 150, g. 
Torymus lissus (Walker) Cresson, 1887 : 237. 
Syntomaspis lissus (Walker) Ashmead, 1887 : 187. 
Callimome aenea Ashmead, 1881a : xxxiii, §, <J. Syn. n. 
Torymus aeneus (Ashmead) Cresson, 1887 : 237. 

Type-material. Callimone lissus Walker. A single male specimen mounted 
on a card is in the BMNH collection, labelled 'B. M. Type Hym. 5. 2287', '1481a' 
and 'Callimome Lissus Walker' by Waterhouse. I designate it LECTOTYPE. 
Described from St John's Bluff, Florida. Although the lectotype is a male, it 
can be placed as the species now called aeneus in the North American literature. 

Callimone aenea Ashmead. This was described from several female and male 
specimens from Jacksonville, Florida; Huber (1927 : 56) designated a lectotype. 
Two other species are at present listed as synonyms of this one, but Torymus is 
now being revised. This restudy might result in some change in the synonymy. 

Distribution. Florida. 

Biology. Emerges from galls of Disholcaspis on white oak. 

Torymus splendidulus Dalla Torre 

Callimome splendidus Walker, 1844 : 14, $. 

Torymus splendidus (Walker) Cresson, 1887 : 237. [Homonym of Torymus splendidus Foerster, 

1841 : 32.] 
Torymus splendidulus Dalla Torre, 1898 : 313. [Replacement name.] 

Type-material. One female specimen mounted on a small pin on a card is 
in the BMNH Collection, labelled 'B. M. Type Hym. 5. 48; St. Martin's Falls' 
and 'Callimome splendidus Walker' by Waterhouse. I designate it LECTOTYPE. 
The specimen is in only fair condition, each antenna being broken off beyond f 4 , 
the thorax distorted by the mounting pin, the ovipositor sheaths lost, but the ovi- 
positor itself remaining. 

Remarks. This species was not recognized by Huber (1927 : 101) and it clearly 
is not included in his revision under some other name. It also is not included in 
the unpublished, but widely circulated, manuscript classification of North American 
Torymus prepared by Lienk (1951). I find no specimens of it in the USNM collection. 



SPECIES OF CHALCIDOIDEA 153 

It has the following key characters. Stigmal vein petiolate; scutellum without 
cross-furrow; antennal scape pale tan, flagellum black; head and thorax dark 
reddish bronze-green; legs, including apices of coxae, bright yellow-tan; gaster 
iridescent green-bronze with a small ventral area at base yellowish; ovipositor 
slightly longer than gaster; propodeum with moderately rough sculpture and a 
prominent median carina that bifurcates near base. 

Distribution. Ontario. 



Torymus theon (Walker) 

Callimome theon Walker, 1843a : 149, 9- 
Syntomaspis theon (Walker) Ashmead, 1887a : 187. 
Torymus theon (Walker) Peck, 195 1 : 526. 

Type-material. There are two female specimens mounted on cards in the 
BMNH collection, one of which is labelled, 'B. M. Type Hym. 5. 20; 1480a St. 
John's Bluff' and 'Callimone Theon Walker' in Waterhouse's hand, and is designated 
LECTOTYPE; this specimen is cemented on its side on the mounting card, with 
the head detached and cemented on the card. 

Remarks. This species cannot at present be placed. It does not run in Huber 
(1927) or in Lienk (1951), and I do not find specimens of it in the USNM collection. 
However, this is a distinctive species that should be possible to place. The antennal 
scape is yellow, the flagellum black; all tibiae are yellow, legs otherwise dark metallic 
green; thorax and gaster dark metallic green with silvery pubescence; stigmal vein 
of forewing subsessile and with three rows of wing bristles radiating from its apex; 
all wing veins yellow; scutellum with a deep cross-furrow, space posterior to this 
furrow shagreened; dorsum of thorax minutely shagreened, with interspersed short 
bristles that arise from obscure pits; propodeum lightly shagreened, almost smooth; 
ovipositor as long as body. The thorax is relatively thickset, with a subacute, 
longitudinal ridge on the posterior third of each lobe of the mesoscutum. 

Distribution. Florida. 



ORMYRIDAE 

Ormyrus labotus Walker 

Ormyrus labotus Walker, 1843a : 148, 9. S- 
Ormyrus andricus Ashmead, 1886 : 128, 9- Syn. n. 

Type-material. Ormyrus labotus Walker. There are one female and two male 
specimens, mounted on cards, in the BMNH collection. The female is labelled 
'B. M. Type Hym. 5. 63; 1479a; St. John's Bluff' and 'Ormyrus Labotus Walker' 
by Waterhouse; I designate it LECTOTYPE. The two males are labelled 'Co-type', 
one '1479b; St. John's Bluff', the other '1479c; St. John's Bluff.' This species is 
clearly the same as andricus Ashmead. 



154 B - D - BURKS 

Ormyrus andricus Ashmead. Described from one female specimen from Jackson- 
ville, Florida. The holotype is USNM no. 41108. 

Distribution. Florida. There are references in the literature to labotus as 
occurring in Kansas and Maryland, but the identifications on which those records 
were based were made without reference to Walker's type. It is not possible 
now to locate the specimens involved, so those records cannot be verified. There 
are many specimens of labotus in the USNM collection, but all are from Florida. 

Biology. Emerges from galls of Callirhytis on white oak. 



PTEROMALIDAE 

Pachyneuron albutius Walker 

Pachynevron albutius Walker, 1843a : 158, $. 

Pachyneuron Albutius Walker, 1846 : 35. [Correction of spelling of generic name.] 

Spalangia syrphi Ashmead, 1881a : 171, §, g. Syn. n. 

Pachyneuron syrphi (Ashmead) Ashmead, 1887a : 193. 

Pachyneuron allograptae Ashmead, 18876 : 15. $, $. Syn. n. 

Type-material. Pachyneuron albutius Walker. There is a single female 
specimen, mounted on a card point, in the BMNH collection, labelled 'B.M. Type 
Hym. 5. 860; 1492a; St. John's Bluff' and 'Pachyneuron Albutius Walker' by 
Waterhouse. I designate it LECTOTYPE. This specimen is in poor condition, 
but it is recognizable as the common Nearctic Pachyneuron parasitic on syrphid 
flies. 

Spalangia syrphi Ashmead. This was originally described from five female 
and two male specimens of which only two female syntypes are now in the USNM 
collection. One female, labelled 'Jacksonville, Fla., Collection Ashmead; Pachy- 
neuron syrphi Ashm.', is designated LECTOTYPE and is USNM Type no. 2823. 

Pachyneuron allograptae Ashmead. This species was described from 'several 
specimens'. There are now five female syntypes in the USNM collection; one, 
labelled 'Jacksonville, Fla., Collection Ashmead; Pachyneuron allograptae Ashm. 
$', is designated LECTOTYPE and is USNM no. 2847. 

Distribution. North America, Central America, West Indies. 

Biology. This species is commonly reared as a primary parasite of syrphid 
flies; see Peck (1963 : 613) for a host list. 

Pteromalus cassotis Walker 

Pteromalus cassotis W T alker, 1847 : 393, §. 

Pteromalus archippi Howard, 1889 : 1872, 1891, $. Syn. n. 

Type-material. Pteromalus cassotis Walker. There are three female specimens, 
on card points, in the BMNH collection, labelled 'B.M. Type Hym. 5. 754a, b, c' 
Each is labelled 'Pteromalus cassotis Walker' by Waterhouse. I designate specimen 



SPECIES OF CHALCIDOIDEA 155 

754c LECTOTYPE. Described from St John's Bluff, Florida. This species is 
the same as the one currently being called archippi. 

Pteromalus archippi Howard. This was described from an unspecified number 
of female specimens from Eastern North America; it is now represented by three 
syntypes in the USNM. One of these, labelled 'Ac. Cat. 815; Ag. Coll. Mich.; 
9-13-87; Pteromalus archippi $ Type How.; U. S. N. M. No. 2675', is designated 
LECTOTYPE. 

Distribution. North America. 

Biology. Most commonly a parasite of nymphalids, but has also been reared 
from pierids and papilionids. 

Neocatolaccus cratylus (Walker) comb. n. 
Pteromalus Cratylus Walker, 1847 : 392, §. 

Type-material. There is a single female specimen mounted on a card point 
in the BMNH collection, labelled 'B.M. Type Hym. 5. 753' and 'Pteromalus Cratylus 
Walker' by Waterhouse. I designate it LECTOTYPE. Described from St John's 
Bluff, Florida. The lectotype is fragmentary, so it probably never can be placed 
specifically, but it is a Neocatolaccus. 

Distribution. Florida. 

Capellia cyrene (Walker) comb. n. 

Micromelus cyrene Walker, 1843a : 154, 0_. 
Boeotomus cyrene (Walker) Cresson, 1887 : 242. 
Callitula cyrene (Walker) Peck, 1951 : 548. 
Pseudomicromelus cyrene (Walker) Burks, 1958 : 76. 

Type-material. There is a single female specimen mounted on a card in the 
BMNH collection, labelled 'B.M. Type Hym. 5. 686; 1487a; St. John's Bluff' and 
'Micromelus Cyrene Walker' by Waterhouse. I designate it LECTOTYPE. This 
specimen was originally cemented on its side on a card in such a way that study 
of it was virtually impossible. Dr Boucek successfully remounted it on a card 
with the wings spread and the dorsal side up, after which delicate operation its 
characters were clearly evident. He and I agree that this belongs in the genus 
Capellia Delucchi, close to, but distinct from, rufiventris (Girault). There is a 
specimen of cyrene in the USNM collection, taken at Jacksonville, Fla., by Ashmead 
many years ago. 

Distribution. Florida. 

Callitula cyrnus (Walker) comb. n. 

Lampretatus cyrnus Walker, 1843a : 157. 
Lamprotatus cyrnus (Walker) Walker, 1846 : 32. 
Cryptoprymnus illinoensis Ashmead, 1896 : 228, °- Syn. n. 
Callitula illinoensis (Ashmead) Peck, 195 1 : 548. 



156 B. D. BURKS 

Type-material. Lampretatus cyrnus Walker. There are eight syntypes moun- 
ted on card points in the BMNH collection, one of which is labelled 'B.M. Type 
Hym. 5. 820; St. John's Bluff' and 'Lamprotatus Cyrnus Walker' by Waterhouse, 
and is designated LECTOTYPE. The other seven specimens, all labelled as 
'Cotypes' of this species, are not congeneric with the lectotype. The lectotype 
is recognizable as the male of the rather rare North American species described by 
Ashmead as illinoensis. 

Cryptoprymnus illinoensis Ashmead. This was described from one female speci- 
men from Algonquin, Illinois. The holotype is USNM no. 26035. 

Distribution. Maryland south to Florida west to Illinois. 

Biology. This species has been reared from the Hessian fly, Mayetiola destructor 
(Say), and other small Dipt era associated with grasses. 



Habrocytus damo (Walker) 

Pteromalus Damo Walker, 1847 : 395, 9- 
Habrocytus damo (Walker) Graham, 1969 : 561. 

Type-material. There is one female specimen mounted on a card point in 
the BMNH collection, labelled 'B.M. Type Hym. 5. 759' and 'Pteromalus Damo 
Walker' by Waterhouse. I designate it LECTOTYPE. Described from St John's 
Bluff, Florida. This lectotype specimen lacks a head, and the legs and wings are 
entangled in the mounting medium. I agree with Graham's generic placement 
of it ; it cannot be placed specifically until the North American species of Habrocytus 
are revised. 

Distribution. Florida. 



Trichomalus deiphon (Walker) comb. n. 

Metopon deiphon Walker, 1843a : 161, 9- 
Psilocera deiphon (Walker) Walker, 1846 : 36. 

Type-material. There is one female specimen mounted on a card point in 
the BMNH collection, labelled 'B.M. Type Hym. 5. 693; 1495a; St. John's Bluff' 
and 'Psilocera Deiphon Walker' by Waterhouse. I designate it LECTOTYPE. 
Graham (1969 : 467) examined this specimen and excluded it from Psilocera and 
was of the opinion that it was near Spaniopus Walker or Trichomalus Thomson. 
I believe it belongs in Trichomalus. There are no specimens of this species in the 
USNM collection. 

Distribution. Florida. 



Seladerma diaeus (Walker) 

Lamprotatus Diaeus Walker, 1844 : 16, $. 
Seladerma diaeus (Walker) Graham, 1969 : 203. 



SPECIES OF CHALCIDOIDEA 157 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 822; 44 17; Martin's Falls; Hudson's Bay; 
Bred from pupa of Vanessa Cardui' and 'Lamprotatus Diaeus Walker' by Water- 
house. Graham (1969 : 203) designated a lectotype for this species and transferred 
it to Seladerma Walker. 

Distribution. Ontario. 

Biology. Graham (1969 : 203) has shown that the host label on the type of 
this species is erroneous and should have referred to Pteromalus puparum (L.) 
instead. His opinion is that the true host of diaeus is probably an agromyzid. 

Pteromalus doryssus Walker 

Pteromalus doryssus Walker, 1847 : 395, Q, $. 

Type-material. Missing. Described from St John's Bluff, Florida. 
Distribution. Florida. 

Zatropis dymnus (Walker) comb. n. 

Pteromalus Dymnus Walker, 1847 : 397, Cj- 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 760' and 'Pteromalus Dymnus Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. This lectotype specimen lacks a head, so its specific placement will prob- 
ably never be possible, but it belongs in the genus Zatropis Crawford. 

Distribution. Florida. 

Norbanus dysaules Walker 

Norbanus dysaules Walker, 1843a : 159, <3\ $. 

Type-material. There are ten female and five male specimens, all on card 
points, in the BMNH collection. They are labelled 'B.M. Type Hym. 5. 694' 
and each bears the Waterhouse labels 'St. John's Bluff; Norbanus Dysaules Walker.' 
The female specimen bearing the number 1493a is designated LECTOTYPE. 
There is an additional female paralectotype in the USNM, received through exchange 
in 1949. 

Remarks. This species, and through it, the genus Norbanus, has long been cor- 
rectly placed in the North American literature. Graham (1969 : 438) examined 
the type-series of dysaules. 

Distribution. Florida. 

Habrocytus epicles (Walker) 

Pteromalus Epicles Walker, 1847 : 394. $. 
Habrocytus epicles (Walker) Graham, 1969 : 561. 



158 B. D. BURKS 

Type-material. There is one female specimen mounted on a card point in the 
BMNH collection, labelled 'B.M. Type Hym. 5. 758' and 'Pteromalus epicles Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. I agree with Graham's generic assignment for this species, but it cannot 
be placed specifically until the North American species of Habrocytus are revised. 

Distribution. Florida. 



Pteromalus eryx Walker 

Pteromalus Eryx Walker, 1847 : 397, <J. 

Type-material. There is one male specimen mounted on a card point in the 
BMNH collection, labelled 'B.M. Type Hym. 5. 761' and 'Pteromalus Eryx Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. This lectotype specimen is in very poor condition, lacking the head and 
anterior legs. It obviously is not Pteromalus, as that genus is currently defined, 
but neither Dr Boucek nor I can place it. I made an exhaustive search through 
the USNM collection, but did not find a pteromalid male that seemed likely to 
be the same as this one. This species should be left unplaced. 

Distribution. Florida. 



Trichomalus eurypon (Walker) comb. n. 

Pteromalus eurypon Walker, 1847 : 398, $. 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 762' and 'Pteromalus Eurypon Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. The single specimen is in poor condition, with all wings and the apex 
of the abdomen missing. It should be placed in the genus Trichomalus Thomson, 
but it is doubtful that it can be placed specifically, because of its fragmentary 
condition. 

Distribution. Florida. 

Pteromalus euthymus Walker 

Pteromalus euthymus Walker, 1847 : 393, g. 

Type-material. Missing. Described from St John's Bluff, Florida. 
Distribution. Florida. 



Trichomalus habis (Walker) comb. n. 

Lampretatus habis Walker, 1843a : 155, Q.. 
Lamprotatus habis (Walker) Walker, 1846 : 32. 



SPECIES OF CHALCIDOIDEA 159 

Type-material. There is one female specimen mounted on a card point in 
the BMNH collection, labelled 'B.M. Type Hym. 5. 818; 1488a; St. John's Bluff' 
and 'Lamprotatus Habis Walker' by Waterhouse. I designate it LECTOTYPE. 
The specimen is almost intact, but the wings are cemented together. This belongs 
in Trichomalus Thomson; I find no specimens of it in the USNM collection. 

Distribution. Florida. 



Zatropis hermeas (Walker) comb. n. 

Pteromalus hermeas Walker, 1847 : 394, $. 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 756' and 'Pteromalus Hermeas Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. This lectotype specimen is badly broken, but the fragments are cemented 
to the card point. It can be placed in the genus Zatropis Crawford, but its specific 
placement is equivocal. It may be the same as the species now being called Z. 
incertus (Ashmead, 1893 : 186). 

Distribution. Florida. 



Acaenacis lausus (Walker) comb. n. 

Pteromalus Lausus Walker, 1847 : 392, $. 
Pteromalus Dipsas Walker, 1847 : 394, <J. Syn. n. 
Semiotellus ficigerae Ashmead, 1885 : xvii, 9. <$■ Syn. n. 
Acaenacis ficigerae (Ashmead) Girault, 1917 : ia. 

Type-material. Pteromalus lausus Walker. There are two female specimens 
on card points in the BMNH collection, labelled 'B.M. Type Hym. 5. 752a, 752b' 
and 'Pteromalus Lausus Walker' by Waterhouse. I designate specimen 752a as 
LECTOTYPE. Described from St John's Bluff, Florida. 

Pteromalus dipsas Walker. There is one male specimen on a card point in the 
BMNH collection, labelled 'B.M. Type Hym. 5. 755' and 'Pteromalus Dipsas Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. This is the male that should be associated with the female described 
as lausus. 

Semiotellus ficigerae Ashmead. This species was described from an unspecified 
number of male and female specimens; there are now four female and three male 
syntypes in the USNM collection. One female labelled 'Jacksonville, Fla., Collec- 
tion Ashmead,' is designated LECTOTYPE. This species agrees in both sexes 
with lausus and dipsas. Girault (1917 : ia) designated ficigerae as type-species 
of his genus Acaenacis. 

Distribution. Florida. 

Biology. Has been reared from a Disholcaspis gall on white oak. 



160 B. D. BURKS 

Mesopolobus oeax (Walker) comb. n. 

Pteromalus oeax Walker, 1847 : 395, 9. 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 763' and 'Pteromalus Oeax Walker' by 
Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. The lone specimen is badly preserved, but it is a species of Mesopolobus 
Westwood. The species of that genus need to be revised for North America. 

Distribution. Florida. 



Zatropis orontas (Walker) comb. n. 

Pteromalus Orontas Walker, 1847 : 396, $. 

Meraporus bruchivorus Ashmead, 1893 : x 6i, $, S- Syn. n. 

Zatropis bruchivorus (Ashmead) Crawford, 1921 : 171. 

Type-material. Pteromalus orontas Walker. There are three female specimens 
on card points in the BMNH collection, labelled 'B.M. Type Hym. 5. 766a, b, c' 
and 'Pteromalus Orontas Walker' by Waterhouse. I designate specimen 766b 
LECTOTYPE. Described from St John's Bluff, Florida. The specimens labelled 
766a and 766c are not congeneric with the lectotype. This lectotype is the same 
as the species now being called Zatropis bruchivorus (Ashmead) in the North 
American literature. 

Meraporus bruchivorus Ashmead. This was described from an unspecified 
number of female and male specimens from localities in Kansas, Indiana, and Iowa; 
there are now 6 $, 4 $ syntypes in the USNM. I designate a female specimen 
labelled 'Nov.; Riley Co., Ks., Marlatt; Meraporus bruchivorus $ Ashm.,' as LECTO- 
TYPE. 

Distribution. Maryland south to Florida, west to Iowa, Kansas, Arizona 
and California; Sonora, Mexico. 

Biology. Has been reared from Acanthoscelides submuticus (Sharp) and other 
bruchids. 

Norbanus pisius Walker 

Norbanus pisius Walker, 1843a : 160, $. 
Pteromalus Hybreas Walker, 1847 : 397, $. Syn. n. 

Type-material. Norbanus pisius Walker. There is one female specimen 
on a card point, with broken parts mounted separately on a card, in the BMNH 
collection; it is labelled 'B.M. Type Hym. 5. 695; 1494a; St. John's Bluff' and 
'Norbanus Pisius Walker' by Waterhouse. I designate it LECTOTYPE. This 
species is correctly identified in the USNM collection. 

Pteromalus hybreas Walker. There is one badly broken female specimen on 
a card point in the BMNH collection, labelled 'B.M. Type Hym. 5. 764' and 
'Pteromalus Hybraeas Walker' by Waterhouse. I designate it LECTOTYPE. 



SPECIES OF CHALCIDOIDEA 161 

Described from St John's Bluff, Florida; it is a Norbanus, and the remaining parts 
all agree with pisius. 

Distribution. Florida. 

Biology. There is a specimen of pisius in the USNM collection from Clewiston, 
Florida, that was reared from Euscepes porcellus Boheman, a sweet potato weevil. 

Lamprotatus salemus (Walker) 

Lampretatus salemus Walker, 1843a : 156, $. 
Lamprotatus Salemus (Walker) Walker, 1846 : 32. 

Type-material. Only a female gaster and one leg (which may not belong 
together) remain in the BMNH collection, labelled 'B.M. Type Hym. 5. 819; 1489a; 
St. John's Bluff' and 'Lamprotatus Salemus Walker' by Waterhouse. I have not 
designated this as lectotype, because a better specimen might eventually be found. 
This species should be unplaced, at least for the present. 

Distribution. Florida. 



Zatropis timaea (Walker) comb. n. 

Pteromalus timaea Walker, 1847 : 396, $. 
Habrocytus timaea (Walker) Peck, 1951 : 563. 

Type-material. There is one female specimen on a card point in the BMNH 
collection, labelled 'B. M. Type Hym. 5. 757' and 'Pteromalus Timaea Walker' 
by Waterhouse. I designate it LECTOTYPE. Described from St John's Bluff, 
Florida. This clearly is a species of Zatropis Crawford. There is in the USNM 
collection a specimen labelled 'Habrocytus timaea Walk.' by A. A. Girault. That 
probably was the basis for the transfer of this species to Habrocytus, but Girault's 
identification was made without reference to Walker's type. 

Distribution. Florida. 



Cyrtogaster trypherus (Walker) comb. n. 

Lampretatus trypherus Walker, 1843a : 158, '§' = <$. 
Lamprotatus Trypherus (Walker) Walker, 1846 : 32. 
Cyrtogaster glasgowi Crawford, 1914 : 36, §. $■ Syn. n. 

Type-material. Lampretatus trypherus Walker. There is one male specimen 
on a card point in the BMNH collection, labelled 'B.M. Type Hym. 5. 821; 1491a; 
St. John's Bluff' and 'Lamprotatus Trypherus Walker' by Waterhouse. I desig- 
nate it LECTOTYPE. This specimen shows the distinctive characters of the male 
of the species currently being identified as Cyrtogaster glasgowi Crawford. It 
has enormous maxillae, the propodeum has lineolate-rugose sculpture, and the legs 
are bright tan in contrast with the dark metallic green body. No other known 
North American male pteromalid is similar. 



i6.z B. D. BURKS 

Cyrtogaster glasgowi Crawford. This was described from two female and one 
male specimens from Urbana, Illinois; the designated female holotype is USNM 
No. 18246. 

Distribution. Delaware south to Florida west to Missouri and California. 

Biology. Has been reared from the pupae of an ephydrid, Brachydeutra argen- 
tata (Walker), and of a Drosophila sp. It also is collected fairly often in alfalfa 
fields. 

EURYTOMIDAE 

Eurytoma abatos Walker 

Eurytoma abatos Walker, 1843a : 152, §. 

Eurytoma crassineura Ashmead, 1894 : 324, $, £. Syn. n. 

Type-material. Eurytoma abatos Walker. One female specimen, mounted 
on a card point, is in the BMNH collection, labelled 'B.M. Type Hym. 5. 579; 
1484a; St. John's Bluff' and 'Eurytoma Abatos Walker' by Waterhouse, also 'abatos 
type' in Walker's hand. I designate it LECTOTYPE. This is a small, light 
coloured specimen of the species currently called Eurytoma crassineura Ashmead 
in the North American literature. I am indebted to Dr Boucek for sending me 
additional information about the type of abatos after I had returned to Washington. 

Eurytoma crassineura Ashmead. Bugbee (1967 : 500) has designated a $ lecto- 
type for this species, which was described from Morgantown, West Virginia. It 
is in the USNM, type no. 25508. 

Distribution. Occurs in southern Canada and throughout the United States. 

Biology. A primary parasite of scolytid beetles. 

Eurytoma cretheis Walker 

Eurytoma cretheis Walker, 1843a : 150, 9. 

Type-material. A single female specimen, on a card point, is in the BMNH 
collection, labelled 'B.M. Type Hym. 5. 583; 1482a; St. John's Bluff' and 'Decatoma 
Cretheis Walker' by Waterhouse, and 'cretheis type' in Walker's hand. I designate 
it LECTOTYPE. The specimen is damaged. The head, antennae, both forewings, 
and one hindwing are missing. This fragmentary type cannot be placed to species. 

Distribution. Florida. 

Eurytoma hecale Walker 

Eurytoma hecale Walker, 1843a : 151, §. 

Eurytoma auriceps Walsh, 1870 : 299, §, <$. Syn. n. 

Type-material. Eurytoma hecale Walker. There are two female specimens 
under this name in the BMNH collection, one of which is mounted on a card point, 



SPECIES OF CHALCIDOIDEA 163 

and labelled 'B.M. Type Hym. 5. 580; 1483a; St. John's Bluff' and 'Eurytoma 
hecale Walker' by Waterhouse, also 'hecale type' in Walker's hand; I designate 
it LECTOTYPE. The other specimen is a different species, so labelled in 1955 
by G. J. Kerrich. The lectotype is recognizably the same as the species currently 
being identified as E. auriceps Walsh in North America. 

Eurytoma auriceps Walsh. The types of this species, described from Illinois, 
were destroyed in the Chicago fire of 1871 (Burks, 1953 : 16). Bugbee (1967 : 479) 
has recharacterized this species in conformity with the current identification of 
it; his material is in the USNM. 

Distribution. Eastern North America. 

Biology. A parasite of cynipid gall makers on white oaks. 



Harmolita iphis (Walker) comb. n. 

Eurytoma iphis Walker, 1846 : 85, 9- 

Type-material. One female specimen, on a card point, is in the BMNH collec- 
tion, labelled 'B.M. Type Hym. 5. 2288' and '1529a; St. John's Bluff' and 'Eurytoma 
Iphis Walker' by Waterhouse. I designate it LECTOTYPE. 

Distribution'. Florida. 

Eurytomocharis pythes (Walker) comb. n. 
Eurytoma pythes Walker, 1843a : 154, Cj- 

Type-material. One female specimen, mounted on a card point, is in the 
BMNH collection, labelled 'B.M. Type Hym. 5. 581; 1486a; St. John's Bluff' and 
'Eurytoma Pythes Walker' by Waterhouse. I designate it LECTOTYPE. 

Distribution. Florida. 

Tenuipetiolus teredon (Walker) comb. n. 

Eurytoma teredon Walker, 1843a : 153, §, <$. 

Eurytoma albipes Ashmead, 1881& : xxxi, 9. S- Syn. n. 

Tenuipetiolus albipes (Ashmead) Bugbee, 1951 : 37. 

Type-material. Eurytoma teredon Walker. There are one female and three 
male specimens, all on card points, in the BMNH collection. I designate the 
female as LECTOTYPE. It is labelled 'B.M. Type Hym. 5. 582' and 'Cotype; 
1485c; St. John's Bluff' and 'Eurytoma Teredon Walker' by Waterhouse. It 
should be noted that there is considerable antigeny in this species, but these quite 
dissimilar males and females are conspecific and were correctly associated by Walker. 
This species is the distinctive one Bugbee (1951 : 37) selected as type-species for 
his genus Tenuipetiolus. 

Eurytoma albipes Ashmead. This species was described from an unspecified 
number of female and male specimens. At present only one broken female specimen 



164 B. D. BURKS 

of it can be found in the USNM collection. I designate it LECTOTYPE. It is 
labelled 'Jacksonville, Fla.; Collection Ashmead; Type no. 2822 U.S.N.M.; Eurytoma 
albipes Ashm.' 

Distribution. New York south to Florida, west to Illinois, Missouri, Texas. 

Biology. A parasite of cecidomyiid and cynipid gall makers. 



CHALCIDIDAE 

Brachymeria alexinus (Walker) comb. n. 

Perilampus Alexinus Walker, 1846 : 89. 

Type-material. There are two male specimens, each mounted on a small 
pin on a card, in the BMNH collection. One bears a label, added long after Walker's 
time, which states in Latin 'True Chalcis, without hind legs — Biro'. This specimen, 
also labelled 'B.M. Type Hym. 5. 2289', is designated LECTOTYPE. The second 
specimen also lacking the hind legs, is labelled 'Brachymeria sp., G. J. Kerrich, 
1958.' Species described from Georgia. 

Remarks. It is obvious that the hind legs, with their enlarged femora, were 
missing from these specimens at the time Walker described alexinus. Had they 
been present, he most certainly would not have placed the species in Perilampus. 
The original description, however, has never sounded very much like that of a 
perilampid; my guess had been that alexinus was some sort of eurytomid. When 
I saw the types, however, I recognized them as a well known Brachymeria secondary 
parasite of grasshoppers, one of Holarctic distribution and one long known in 
the world literature as Brachymeria color adensis (Cresson, 1872). If necessary, 
I shall request the International Commission to suppress Walker's name for this 
species in favour of Cresson's name. The name coloradensis has been used in manuals 
or revisions published in Japan (Habu, i960 : 159; 1962 : 27), Russia (Nikolskaya, 
i960 : 85), Czechoslovakia (Boucek, 1957 : 210), France (Steffan, 1959 : 42) and 
North America (Burks, i960 : 250). 



Haltichella onatas (Walker) 

Hockevia onatas Walker, 1843a : 146, ?. 

Haltichella onatas (Walker) Walker, 1846 : 7. 

Conura Onatas (Walker) Walker, 1871 : 41. 

Haltichella longicovnis Ashmead, 1887a : 185, $. Syn. n. 

Type-material. Hockeria onatas Walker. There are three female specimens, 
mounted on card points, in the BMNH collection. One is labelled 'B.M. Type 
Hym. 5. 553; 1477a; St. John's Bluff' and 'Hockeria Onatas Walker' by Waterhouse, 
and I designate it LECTOTYPE. One of the other two, labelled '1477b' is frag- 
mentary; the other, labelled '14776', is virtually intact. This is the species currently 
being identified in North America as Haltichella longicornis Ashmead. 



SPECIES OF CHALCIDOIDEA 165 

Haltichella longicornis Ashmead. Described from one female specimen from 
Jacksonville, Florida, USNM Holotype no. 20666. 

Distribution. South-eastern United States. 

Biology. A primary parasite of moths of the families Oecophoridae and Gele- 
chiidae. 

Spilochalcis side (Walker) 

Smiera side Walker, 1843a : 145, £• 
Smicra torvina Cresson, 1872 : 40. ty. 
Spilochalcis side (Walker) Burks, 1940 : 336. 

Type-material. One female specimen, mounted on a card, is in the BMNH 
collection, labelled 'B.M. Hym. 5. 481; 1476a; St. John's Bluff' and 'Smicra Side 
Walker' by Waterhouse, also 'Side' in Walker's hand. I designate it LECTOTYPE. 

Distribution. Throughout southern Canada and United States. 

Biology. Has a lengthy host list; see Peck (1963 : 888). 

Spilochalcis transitiva (Walker) 

Smiera transitiva Walker, 1862 : 371, $. 
Spilochalcis transitiva (Walker) Howard, 1897 : 130. 

Type-material. One female specimen mounted on a small pin on a card is in 
the BMNH collection. It is labelled 'B.M. Type Hym. 5. 537; 43-15; North America' 
and 'Smiera transitiva Walker' by Waterhouse, also 'Tetrasmicra transitiva' 
in an unknown hand. I designate it LECTOTYPE. The current application of 
the name is correct. Described from Eastern Florida. 

Distribution. Georgia, Florida, Texas, Arizona; Mexico, Central America; 
Cuba, Grenada, Jamaica. 

Biology. A primary parasite of the pierid butterfly, Catopsilia eubule (L.). 

Haltichella xanticles (Walker) 

Hockeria xanticles Walker, 1843a : 147, £• 
Haltichella xanticles (Walker) Walker, 1846 : 7. 
Conura Xanticles (Walker) Walker, 1871 : 41. 
Haltichella americana Howard, 1885 : 36, 9. Syn. n. 

Type-material. Hockeria xanticles Walker. There are three female specimens 
on card points in the BMNH collection. One is labelled 'B.M. Type Hym. 5. 554; 
1478a; St. John's Bluff' and 'Hockeria Xanticles Walker' by Waterhouse. Another 
is labelled, 'Cotype; 1478b'; the third is labelled 'Cotype; 1478c' The specimen 
labelled '1478c' is designated LECTOTYPE. This species is being correctly identi- 
fied at present in North American literature, and Howard's americana is the same. 

Haltichella americana Howard. Described from two female syntypes which 
are, unfortunately, not conspecific. One, labelled 'Washington, D. C, 2. 7; Type 



166 B. D. BURKS 

No. 2627 U. S. N. M.; Haltichella americana How. ms.', is the one on which the 
name americana has always been based. I designate it LECTOTYPE. The 
other syntype has the same labels, plus a 'Schwarz Coll.' label. It is a specimen 
of Haltichella onatas (Walker). 

Distribution. Quebec south to Florida west to Saskatchewan and Texas. 

Biology. This is a primary parasite of small moths of the families Olethreutidae, 
Coleophoridae, Lyonetiidae. It also has been reared as a secondary parasite, 
emerging from the cocoons of Apanteles. 



LEUCOSPIDIDAE 

Leucospis affinis Say 

Leucospis ujfinis Say, 1824 : 326, 9. cj- 

Leucospis Canadensis Walker, i860 : 17, £. [Synonymized by Cresson, 1872 : 32.] 

Type-material. Leucospis affinis Say. Type lost, but the identity of this 
species has long been accepted in North American literature on the basis of the 
original description and the redescription of Cresson (1872 : 32). 

Leucospis canadensis Walker. There is one male specimen mounted on a small 
pin on a card in the BMNH collection. It is labelled 'B.M. Type Hym. 5. 91; 59. 
13 <$', Canada; Pres. by W. S. M. D'Urban; 75 C. H.; 56. 1859-130' and 'Leucospis 
canadensis Walker' by Waterhouse, also 'Leucospis affinis Say <§, J. Waterston 
det.' I designate it LECTOTYPE. Both antennae are broken off beyond the 
pedicel, but otherwise this is an intact male specimen of the form currently being 
identified as affinis affinis Say in eastern North America. There is a Floridian 
subspecies that differs. 



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INDEX 
Synonyms are in italics 



abatos, 162 
Acaenacis, 159 
Achrysocharella, 142 
Acrias, 146 
aea, 151 
aenea, 152 
affinis, 166 
albifrons, 151 
albipes, 163 
albutius, 154 
alexinus, 164 
allograptae, 154 
amelanchieris, 151 
americana, 165 
americanum, 147 
andricus, 153 
Aprostocetus, 142 
avchippi, 154 
Aulogymnus, 146 
auriceps, 162 

bolus, 147 
Brachymeria, 164 
bruchivorus, 160 



calavius, 141 
Callitula, 155 
canadensis, 166 
Capellia, 155 
cassotis, 154 
cecidomyiae, 151 
Chalcididae, 164 
Chrysopophagus, 147 
Cirrospilus, 143 
columbiana, 144 
cvassineura, 162 
cratylus, 155 
cretheis, 162 
cyrene, 155 
cyriades, 142 
cyrnus, 155 
Cyrtogaster, 161 



damastes, 142 
damo, 156 
deiphon, 156 
diaeus, 156 
dipsas, 159 
doryssus, 157 
dymnus, 157 
dysaules, 157 

Encyrtidae, 147 
Encyrtus, 148 
entellus, 150 
epicaste, 149 
epicles, 157 
epidius, 142 
eryx, 158 
Euderus, 144 
Eulophidae, 141 
eunapius, 143 
Eupelmella, 149 
Eupelmidae, 149 
Eupelmus, 149, 150 
eurypon, 158 
Eurytoma, 162 
Eurytomidae, 162 
Eurytomocharis, 163 
euthymus, 158 

ficigerae, 159 
fiaccus, 147 
flavipes, 147 
fonteia, 149 

gala, 143 

Galeopsomyia, 144 

gastron, 148 

glasgowi, 161 

gobryas, 145 

granulatus, Aprostocetus, 142 

granulatus, Tetrastichus, 143 

habis, 158 
Habrocytus, 156, 157 



170 



B. D. BURKS 



haemon, 144 
Haltichella, 164, 165 
Harmolita, 163 
hecale, 162 
herillus, 144 
hermeas, 159 
Horismenus, 147 
hyalinus, 150 
hybreas, 160 

illinoensis , 155 
imbrasus, 145 
iphinoe, 145 
iphis, 163 

labotus, 153 
lamachus, 150 
Lamprotatus, 161 
lausus, 159 
Leipocrossus, 144 
lepreos, 150 
Leucospididae, 166 
Leucospis, 166 
levana, 145 
lissus, 152 
longicornis, 164 

Megastigmus, 151 
Mesopolobus, 160 
meteori, 149 
minio, 145 
minyas, 146 

Necremnus, 142, 145 
Neocatolaccus, 155 
Neochrysocharis, 145 
nileus, 146 
Norbanus, 157, 160 
Notanisomorpha, 14 1 

oeax, 160 
onatas, 164 
Ormyridae, 153 



Ormyrus, 153 
orontas, 160 

Pa chy neuron, 154 
Perilampidae, 150 
Perilampus, 150 
pisius, 160 
Pnigalio, 145 
proximus, 145 
Pseudencyrtus, 147 
Psyllaephagus, 148 
Pteromalidae, 154 
Pteromalus, 154, 157, 158 
pythes, 163 

reate, 148 

salemus, 161 
sardus, 147 
Seladerma, 156 
side, 165 
solus, 148 
Spilochalcis, 165 
splendidulus, 152 
splendidus, 152 
Stenomesius, 145 
strobiloides , 151 
syyphi, 154 

Tenuipetiolus, 163 
teredon, 163 
Tetrastichus, 143 
theon, 153 
timaea, 161 
torvina, 165 
Torymidae, 151 
Torymus, 151, 152, 153 
transitiva, 165 
Trichomalus, 156, 158 
trypherus, 161 

vectius, 148 

xanticles, 165 

Zatropis, 157, 159, 160, 161 



B. D. Burks, B.A., M.A., Ph.D. 
Systematic Entomology Laboratory 
Agricultural Research Service, U.S.D.A. 
Washington, D.C. 20250 
U.S.A. 



ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177: 
18 plates, 270 text-figures. August 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 
1965. £3.25. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text -figures. January, 1967. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae) . Pp. 119: 14 plates, 146 
text -figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

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82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£i9- 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text -figures. 
February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text-figures. October, 1973. £12.30. 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text -figures. December, 1973. £9-55- 

printed by Unwin Brothers Limited the gresham press old woking surrey England 



u» \ 






AN ILLUSTRATED LIST OF THEx. 

TYPE-SPECIMENS OF THE 

HELICONIINAE 

(LEPIDOPTERA : NYMPHALIDAE) 

IN THE BRITISH MUSEUM 

(NATURAL HISTORY) 



P. R. ACKERY 

AND 

R. L. SMILES 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 5 

LONDON : 1976 



AN ILLUSTRATED LIST OF THE 

TYPE-SPECIMENS OF THE 

HELICONIINAE (LEPIDOPTERA : NYMPHALIDAE) 

IN THE BRITISH MUSEUM (NATURAL HISTORY) 



BY 

PHILLIP RONALD ACKERY 

v 

AND 

ROBERT LESLIE SMILES 



Pp. 171-214; 39 Plates 



BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 5 

LONDON : 1976 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in Jive series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 5 of the Entomology 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Ent.). 



ISSN 0524-6431 



Trustees of the British Museum (Natural History), 1976 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued January, 1976 Price £10.70 




AN ILLUSTRATED LIST OF THE 

TYPE-SPECIMENS OF THE W^* 

HELICONIINAE (LEPIDOPTERA : NYMPHALIDAE) /* 
IN THE BRITISH MUSEUM (NATURAL HISTORY) 

By P. R. ACKERY & R. L. SMILES 

CONTENTS 

Page 

Synopsis ........... 173 

Introduction .......... 173 

A List of the type-specimens of the Heliconiinae 

in the British Museum (Natural History) .... 176 

References . . . . . . . . . . . 210 

SYNOPSIS 

A list is given of the type-specimens of the 401 nominal species, subspecies and infrasubspecific 
forms, currently placed in the subfamily Heliconiinae, represented by type-material in the 
British Museum (Natural History). A specimen from the type-series of each taxon is illustrated 
at two-thirds natural size, the figures being arranged systematically. 



INTRODUCTION 

The Heliconiinae comprise some seventy species, but such is the extreme variation 
in pattern within many of them that a multitude of varietal forms have been 
described; about half the names involved are represented by type-material in the 
British Museum (Natural History) (BMNH). Some of the species show quite 
exceptional variation, although individual patterns are often constant within large, 
well-defined geographical areas. However, where these forms meet there are areas 
of transition, sometimes less than 50 kilometres wide (Turner, 1971), in which 
there is extensive mixing of pattern elements. It was often from these regions, 
usually the more accessible parts of tropical South America, that collectors sent 
material in the past. Many of the forms described by Joicey & Kaye (1917; 1919) 
originated from such a transitional zone in French Guiana. 

Early authors during the nineteenth century usually described these transitional 
forms and subspecies as distinct species. The most notable attempts at a ration- 
alization of this situation are to be found in the works of Stichel & Riffarth (1905) 
and Eltringham (1916). More recently Emsley (1963; 1964; 1965) completed his 
studies which have provided the basis of much of the current work being carried 
out on the heliconiines. 

Some outline of the principal authors whose type-material is deposited in the 
BMHN is probably useful. As might be expected, types of such authors as Bates, 



174 P- R- ACKERY & R. L. SMILES 

Butler, Hewitson and Joicey are to be found, but in addition, perhaps unexpectedly, 
type-specimens of many major continental authors are present. A brief outline is 
given below of how some of these specimens came to be housed in the BMNH 
collections. 

Fabricius, J. C. (1745-1808). 

There is only one heliconiine type-specimen of Fabricius to be found in the 
BMNH; that is a single male syntype of Papilio phyllis which is housed in the 
collection of Sir Joseph Banks (1743-1820). This collection came to the museum 
from the Linnean Society and is still retained as a separate entity, none of the 
material having been incorporated in the main series. 

Cramer, P. ( -1779) and Stoll, C. ( -1795). 

The question of the authenticity of the type-material of these authors is dealt 
with by Vane-Wright (1975). The probable heliconiine types are here figured on 
a separate plate (PI. 39) and although none agree perfectly with the original 
illustrations of Cramer and Stoll, there seems to be little doubt that their paintings 
are considerably stylized. Certainly the material is contemporary, the names of 
the collectors, van Lennep and van der Capellen, appearing in an 'Acknowledgment 
List' in Cramer's De Uitlandsche Kapellen. The material passed through the 
Felder collection and the Rothschild Bequest to the BMNH collection. 

Felder, C. (1814-1894) and Felder, R. (1842-1871). 

Many of the taxa described by these authors are based in part or in entirety on 
specimens to be found in the Felder collection. However, sometimes the type- 
material is said to be housed in the 'Musei Caes. Viennes'. The latter is now in the 
Naturhistorisches Museum, Vienna, but the Felder collection was obtained by 
Lord Lionel Walter Rothschild and passed to the BMNH through the Rothschild 
Bequest. 

Oberthur, C. (1845-1924) and Boisduval, J. B. A. D. (1799-1879). 

The type-specimens of the forms described by both these authors were normally 
to be found in their own collections. That of Boisduval was obtained by Charles 
Oberthur and when his collection was sold most of the heliconiine butterflies with 
the exception of Heliconius were purchased by the BMNH. The Heliconius passed 
initially into the collection of Mr John Levick, but after his death also came to the 
BMNH. 

Neustetter, H. ( -1958). 

Type-specimens of many forms described by Neustetter were housed in the 
collection of J. J. Joicey and passed into the BMNH collections through the Joicey 
Bequest. Details of further Neustetter types are included by Holzinger & 



TYPE-SPECIMENS OF HELICONIINAE i 75 

Holzinger (1974) in their catalogue of the Heliconiine type-specimens deposited in 
the Naturhistorisches Museum, Vienna (NM, Vienna). 

RlFFARTH, H. (1860-I908). 

The Heliconius collection of Herr Heinrich Riffarth was acquired by Mr J. J. 
Joicey in 1919 and was obtained by the BMNH in the Joicey Bequest. All the 
type-specimens normally bear a characteristic pink 'Original' type label. 

Niepelt, W. (1862-1936). 

Niepelt was an insect dealer and apparently sold many type-specimens to various 
collectors. The major sources of his material in the Museum collections are the 
Adams Bequest, Joicey Bequest and Rothschild Bequest. Many of his type- 
specimens bear 'Original' labels of identical design to those found on Riffarth 
specimens, but others, apparently equally authentic, do not. Perhaps some of 
his type-specimens should only tentatively be considered valid. Certainly there 
are instances where the commercial zeal of some dealers outstripped their taxonomic 
integrity ! 

Stichel, H. (1862-1936). 

The two main sources of Stichel type-material are the Joicey Bequest and the 
Rothschild Bequest. The former appears to have originated from the Riffarth 
collection and generally each specimen bears an 'Original' label of the usual Riffarth 
pattern whereas those from the Rothschild Bequest normally carry a characteristic 
dark red 'Typus' label. 

Kaye, W. J. (1875-1967). 

Although the museum collection is rich in type-material of the forms described 
jointly by Joicey and Kaye, there are very few authentic Kaye types. Rarely the 
occasional specimen has been found, but the great bulk of the Kaye collection was 
purchased by the Allyn Museum of Entomology, Sarasota, Florida. 

In this present work the taxa are listed alphabetically, with a reference to the 
original description together with the original status and combination. The type 
category and the number of type-specimens are given along with the data quoted 
directly from the labels beneath the specimens, the extent of each label being 
indicated by the sign '/'. Metric equivalents are included where necessary for linear 
measurements quoted from data labels. In addition to the locality information, any 
data pertaining to the history of the specimens are given. Such supplementary labels 
are often useful when the authenticity of the type-material may be open to doubt. 
In each case the current status and combination reflects the opinion of Professor K. S. 
Brown (pers. comm., 1974) which, in some cases, is not that of the authors (e.g. Vane- 
Wright, Ackery & Smiles, 1975). Where the data on the labels do not appear to 
correspond with the published data, the latter are also quoted. A reference is 
given to any noted lectotype designation. 



176 P. R. ACKERY & R. L. SMILES 

The preparation of this work has been greatly assisted by Professor Keith S. 
Brown Jr, Sao Paulo, who studied the Heliconiinae collection at the BMNH during 
a visit sponsored by the Conselho Nacional de Pesquisas, Brazil and the Royal 
Society (London). Professor Brown has generously advised the authors during 
all stages of the final preparation of the manuscript. We also wish to thank 
Mr R. I. Vane-Wright for his help and encouragement during the preparation of 
this work. For the photographs, we are grateful to Mr P. V. York, who undertook 
the arduous task of photographing the specimens, and to Mr P. J. Green for his 
willing cooperation. 



A LIST OF THE TYPE-SPECIMENS OF THE HELICONIINAE 
IN THE BRITISH MUSEUM (NATURAL HISTORY) 

acacetes Hewitson, 18696 : 22 (as sp. of Eueides). 3 <J, 4 $ syntypes. Ecuador. Hewitson 
Coll. 79. 69. [PI. 3, fig. 34.] 

Currently placed as a subspecies of Eueides lampeto Bates. 

adonis Riffarth, 19076 : 507 (as form of Heliconius batesi plesseni). 2 $, 1 $ syntypes. 
Original. 1 $, Pastaza sup., Ecuador, Oct-Dez. 06. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. 1 (J, Rothschild Bequest, B.M. 1939-1. 1 $, Ex. Coll. Riffarth. / 
Joicey Bequest. Brit. Mus. 1934-120. [PI. 19, fig. 197.] 

Currently placed as a transitional form between Heliconius melpomene plesseni Riffarth 
and Heliconius melpomene aglaope Felder & Felder. 

adusta Stichel, 1903 : 11 (as ab. of Eueides cleobaea zorcaon). 1 $, 2 $ syntypes. 1 $, Panama, 
Bugaba, e. c. H. Stichel. / Typus / Rothschild Bequest. B. M. 1939-1. 1 $, Honduras. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <j>, Mexico, Vera 
Cruz, e. c. H. Stichel. / Typus. [P. 5, fig. 48.] 

Currently placed as a form of Eueides isabella eva Fabricius. 

Described from Chiriqui, Honduras. 

aerotome Felder & Felder, 1862a : 79 (as sp. of Heliconius). 1 o* syntype. Rio Negro. / 
Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 17, fig. 172.] 
Currently placed as a subspecies of Heliconius ethilla Godart. 

aglaope Felder & Felder, 1862a : 79 (as sp. of Heliconius). 2 6* syntypes. Rio Negro. / 
Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 20, fig. 206.] 
Currently placed as a subspecies of Heliconius melpomene (L.). 

aglaopeia Staudinger, 1896 : 305 (as var. of Heliconius thelxiope). 1 <J syntype. Cayenne. / 
Origin. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 232.] 
Currently placed as a form of Heliconius melpomene thelxiope (Hiibner). 

agnata Stichel, 1906a : 46 (as form of Heliconius erato estrella). Holotype $. Ucayali. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, fig. 300.] 

Currently placed as a transitional form between Heliconius erato microclea Kaye and 
Heliconius erato emma Riffarth. 

aida Neustetter, 1926a : 37 (as form of Heliconius penelope). 1 <£, 1 $ syntypes. Bolivia. 
1925. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 24, fig. 246.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 



TYPE-SPECIMENS OF HELICONIINAE 177 

alba Riffarth, 1900 : 208 (as ab. of Heliconius antiochus). 1 o*» 3 ? syntypes. Original. / Ex. 
Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 g, 1 $, Amazonas Sup. 1 $, 
Manicore. 1 $, Surinam. Coll. Fruhst. [PI. 37, fig. 381.] 

Currently placed as a subspecies of Heliconius antiochus (L.). 

Described from Sarayacu, Amazon sup. Surinam. 

albidior Neustetter, 1928^ : 259 (as form of Heliconius cydno cydnides). Holotype o*. Rio 
Putamayo, Montegoa? / Joicey Bequest. Brit. Mus. 1934-120. [PI. 27, fig. 272.] 

Currently placed as a transitional form between Heliconius cydno cydnides Staudinger 
and Heliconius cydno zelinde Butler. 

albinea Riffarth, 1899 : 408 (as ab. of Heliconius rhea). Holotype $. Surinam (Stichel). / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 37, fig. 379.] 
Currently placed as a form of Heliconius sara thamar (Hiibner). 
Described from a single male specimen only; however, this female is apparently the type. 

albipunctata Riffarth, 1900 : 199 (as ab. of Heliconius zuleika). 3 o\ J ? syntypes. Chiriqui. 
coll. Staud. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 14, 
fig. 142.] 

Currently placed as a form of Heliconius hecale zuleika Hewitson. 

albucilla Bates, 1866 : 88 (as sp. of Heliconius). 2 <J syntypes. Lion Hill, Panama. 
McLeannan. / Godman-Salvin Coll. 1913-2. [PI. 14, fig. 148.] 

Currently placed as a transitional form between Heliconius hecale melicerta Bates and 
Heliconius hecale zuleika Hewitson. 

alexander Neustetter, 1928a : 443 (as form of Heliconius humboldti). 1 $ syntype. Rio 
Itaya by Iquitos. / Type. O. Bang-Haas. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 15, 

fig- I57-] 

Currently placed as a form of Heliconius hecale humboldti Neustetter. 

alithea Hewitson, 18696 : 10 (as sp. of Heliconia). 3 o*. 1 ? syntypes. Ecuador. Hewitson 
Coll. 79. 69. [PI. 28, fig. 282.] 

Currently placed as a subspecies of Heliconius cydno Doubleday. 

amalfreda Riffarth, 1900 : 212 (as ab. of Heliconius phyllis). 2 6* syntypes. Ex. Coll. 
Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q". Man. H. / Original. 1 6*. Berg en 
Daal, Sur., Juli. Michaelis. [PI. 32, fig. 322.] 

Currently placed as a subspecies of Heliconius erato (L.). 

amandoides Neustetter, 1926a : 39 (as form of Heliconius penelope). 1 o* syntype. Bolivia. 
1925. W. Niepelt, Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PL 25, fig. 255.] 
Currently placed as a form of Heliconius melpomene amandus Grose-Smith & Kirby. 

amandus Grose-Smith & Kirby, 1892 : pi. 1, fig. 3 (as sp. of Heliconius). 1 o* syntype. 
Boliv. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 252.] 
Currently placed as a subspecies of Heliconius melpomene (L.). 

amaryllis Felder & Felder, 1862a : 80 (as sp. of Heliconius). 1 6" syntype. Rio Negro. / 
Rothschild Bequest. B. M. 1939-1. [PI. 19, fig. 191.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 

amathusia Cramer, [1777] : 124, pi. 177, fig. F (as sp. of Papilio). 1 6* syntype. No. 6. 
Amathusia. Cr., II, 177, F. / Felder Colin. / Rothschild Bequest. B. M. 1939-L [PI- 39. 
fig- 396.] 

Currently placed as a form of Heliconius doris doris (L.). 

amneris Neustetter, 1926a : 37 (as form of Heliconius penelope). 2 q* syntypes. Bolivia. 1925. 
W. Niepelt, Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 250.] 



178 P. R. ACKERY & R. L. SMILES 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 

atnoena Stichel, 1903 : 13 (as ab. of Eueides lampeto). 1 o*. I $ syntypes. Original. / Ex. 
Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q\ Santa Jnez (Ecuad.), R. Haensch 
S. 1 $, Bolivien. [PL 4, fig. 35.] 

Currently placed as a form of Eueides lampeto acacetes Hewitson. 

amphitrite Riffarth, 1901 : 157 (as subsp. of Heliconius phyllis). 4 6* syntypes. Hillapani, 
Peru. Garlepp. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 33, fig. 337.] 

Currently placed as a subspecies of Heliconius erato (L.). 

anacreon Grose-Smith & Kirby, 1892 : pi. 1, fig. 5 (as sp. of Heliconius). 1 q*. syntype. 
Boliv. / Ex. Grose Smith, 1910. / Joicey Bequest. Brit. Mus. 1934-120. [Pi. 34, fig. 347.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius). 

anacreonides Neustetter, 1925 : 60 (as form of Heliconius phyllis kruegeri). Holotype <J. 
Rio Grande, Bolivia. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 34, fig. 346.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius). 

anactorie Doubleday, 1847 : 103, pi. 15, fig. 4 (as sp. of Heliconia). 1 o* syntype. Bolivia. 
Bridges. 46-76. [PI. 33, fig. 341.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius) . 

anaitis Riffarth, 19076 : 513 (as form of Heliconius erato anacreon). Holotype $. Prov. Sara, 
Dep. St. Cruz, Slid Bolivia. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. [PI- 34- fi g- 35i-] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius). 

anderida Hewitson, [1853] : pi. 1, fig. 2 (as sp. of Heliconia). 1 q" syntype. Caracas. 
Hewitson Coll. 79. 69. [PI. 15, fig. 151.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius). 
Described from Honduras. 

andicola Bates, 18646 : 187 (as sp. of Agraulis). 2 o" syntypes. Godman-Salvin Coll. 1915-3. 
1 o\ Tacunga, Ecuador. Bates Coll. 1 o". Canelos, Ecuador, Bates Coll. [PI. 1, fig. 7.] 
Currently placed as a subspecies of Dione juno (Cramer) . 

andremona Cramer, [1780] : 16, pi. 297, fig. A (as sp. of Papilio). 1 o* syntype. Surinam. 
Coll. Klukenberg. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 39, fig. 393.] 

Currently placed as ^ transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara (Hewitson). 

androdaixa Seitz, 1913 : 393 (as ab. of Heliconius phyllis andremona). Holotype o*- Bragance, 
(Para). M. de Mathan. / Levick Bequest. 1941-83. [PI. 33, fig. 335.] 

Currently placed as a transitional form between Heliconius erato amazona Staudinger and 
Heliconius erato hydara Hewitson. 

antigona Riffarth, 1900 : 210 (as var. of Heliconius hydara). Holotype o*- Columbien. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, Fig. 293.] 
Currently placed as a form of Heliconius erato demophoon Menetries. 
Described from Medellin, Rio Dagua. 



TYPE-SPECIMENS OF HELICONIINAE 179 

aquilionaris Brown, 1975 : 222 (as subsp. of Heliconius congener). Holotype 6*, 5 6*. 5 $ 
paratypes. q* holotype, 1 q\ 3 $ paratypes, Medina, Ost. Colombia, 1500 m. Coll. Fassl. / 
Joicey Bequest. Brit. Mus. 1934-120. 1 q", 1 9 paratypes, Medina, Ost. Colombia, 1500 m. 
Coll. Fassl. / Rothschild Bequest. B. M. 1939-1. 1 o* paratype, Colombie, Env. Bogota. / 
Joicey Bequest. Brit. Mus. 1934-120. 2 (J, 1 $ paratypes, Colombie, Env. Bogota. Frere 
Apollinaire-Marie. 1918. / Levick Bequest. 11941-83. [PI. 37, fig. 384.] 

araguaia Brown, 1975 : 216 (as subsp. of Heliconius wallacei). Holotype S> 7 S paratypes. <$ 
holotype, 3 o* paratypes, R. Araguaya, Prov. Goyaz. June, 1906. (G. A. Baer.) / Rothschild 
Bequest. B. M. 1939-1. 2 $ paratypes, Tocantins. / Ex. Grose Smith. 1910 / Joicey 
Bequest. Brit. Mus. 1934-120. 2 $ paratypes, Araguaya R. E. Reynolds. 90-58. [PI. 8, 
fig. 86.] 

arcuella Druce, 1874 : 156 (as sp. oiHeliconia). 1 o* syntype. Nauta, Ecuador. E. Bartlett. / 
Godman-Salvin Coll. 1913-2. [PI. 10, fig. no.] 

Currently placed as a subspecies of Heliconius numata (Cramer). 

aristiona Hewitson, [1853] : pi. 1, fig. 4 (as sp. of Heliconia). 1 <$ syntype. Colombia, 
Hewitson Coll. 79. 69. [PI. 10, fig. 106.] 

Currently placed as a subspecies of Heliconius numata (Cramer). 

aristomache Riffarth, 1901 : 131 (as subsp. of Heliconius eralo). 3 $, 1 $ syntypes. Original./ 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 3 o* syntypes, Balzapamba, 
(Ecuad.) R. Haensch. S. 1 $ syntype, Palmar, (Ecuad.) R. Haensch. S. [PI. 9, fig. 96.] 
Currently placed as a form of Heliconius doris eratonius Staudinger. 

arquata Stichel, 1903 : 9, pi. 1, fig. 9 (as subsp. of Eueides isabella). 1 q" syntype. S. America, 
Caucathal. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 5, fig. 50.] 
Currently placed as a form of Eueides isabella huebneri Menetries. 

artemis Riffarth, 19076 : 502, pi. 5, fig. 4 (as subsp. of Heliconius novatus). Holotype <J- 
Ostabh. d. Cordillere a. Titicaca S. Bolivia. / Original' / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. n, fig. 118.] 

Currently placed as a form of Heliconius numata leopardus Weymer. 

artifex Stichel, 1899a : 29 (as ab. of Heliconius erato). 2 q" syntypes. 1 o* syntype, Brazilia, 
Espirito Santo, e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. 1 o" syntype, 
Paraguay, H. Stichel. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 35, fig. 353.] 

Currently placed as a form of Heliconius erato phyllis (Fabricius). 

athalia Neustetter, 19276 : 228 (as form of Heliconius melpomene). Holotype o*- St Laurent, 
Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 236.] 

Currently placed as a form of Heliconius melpomene thelxiope (Hiibner). 

The original description is clearly based on one specimen only and the above specimen 
appears to be the true holotype. Holzinger & Holzinger (1974) also doubtfully include a 
specimen in the NM, Vienna as the 'Holotype' of athalia. 

atrosecta Riffarth, 1900 : 202 (as ab. of Heliconius melpomene). 2 o\ 1 9 syntypes. Obidos 
(Michaelis). 1894 / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 22, fig 219.] 

Currently placed as a form of Heliconius melpomene melpomene (L.). 

Described from three males only; however, these specimens apparently constitute the 
syntypic series. 

atthis Doubleday, 1847 : 102, pi. 14, fig. 3 (as sp. of Heliconia). 1 <$ syntype. Guayaquil. 
Dr J. Hooker. 45-142. [PI. 17, fig. 174.] 

Currently placed as a species of Heliconius Kluk. 



180 P. R. ACKERY & R. L. SMILES 

augusta Riffarth, 1900 : 204 (as var. of Heliconius melpomene). 1 <J paratype. Cayenne. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 237.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene thelxiope (Hubner). 

aurelia Neustetter, 19266 : 289 (as form of Heliconius melpomene). 1 o* syntype. Guyane 
franc, Maroni. Le Moult. 1899-1901. / Levick Bequest 1941-83. [PI. 22, fig. 224.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene meriana Turner. 

aurofasciata Neustetter, 19286 : 76 (as form of Heliconius melpomene aglaope). Holotype <J. 
S. Columbien, Mocoa. November 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 21, 
fig- 209.] 

Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene bellula Stichel. 

aurora Bates, 1862 : 555 (as sp. of Heliconius). 1 $ syntype. S. Paulo, Amazons, Bates. / 
Godman-Salvin Coll. 1913-2. [PI. 10, fig. in.] 

Currently placed as a subspecies of Heliconius numata (Cramer). 

australis Brown, 1975 : 218 (as subsp. of Heliconius hecale). 1 o* paratype. Quevedo, W. 
Ecuador. (V. Buchwald). / Rothschild Bequest. B. M. 1939-1. [P. 15, fig. 150.] 

automatia Oberthiir, 1925 : 81, fig. 5 (as sp. of Heliconia). Holotype q\ Guyane Francaise, 
St Laurent du Maroni. Ex. Antonin Garre. 1923. [PL 36,..fig. 368.] 
Currently placed as a synonym of Heliconius demeter bouqueti Noldner. 

aventina Oberthiir, 1925 : 81, fig. 6 (as sp. of Heliconia). Holotype $. Nouve Grenade. / 
Levick Bequest B. M. 1941-83. [PI. 26, fig. 267.] 

Currently placed as a hybrid between Heliconius cydno hermogenes Hewitson and Heliconius 
melpomene melpomene (L.). 

azteka Neustetter, 1928^ : 259 (as form of Heliconius cydno cydnides). Holotype o*. Solano, 
W. Col. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 26, fig. 265.] 

Currently placed as a transitional form between Heliconius cydno cydnides Staudinger and 
Heliconius cydno zelinde Butler. 

barcanti Brown, 1975 : 219 (as subsp. of Heliconius hecale). Holotype o*. 2 o paratypes. 
Carupano, Dec. 1891. C. W. Ellacombe. / Rothschild Bequest. B. M. 1939-1. [PI. 16, 
fig. 160.] 

bari Oberthiir, 1902 : 23, pi. 21, figs 129, 130 (as sp. of Heliconia). 1 o\ 1 9 syntypes. Levick 
Bequest. 1941-83. 1 q* syntype, Guyane Francaise. Coll. C. Bar. 1 $ syntype, Cayenne. 
[PI. 17, fig. 176.] 

Currently placed as a subspecies of Heliconius elevatus Noldner. 

bartletti Druce, 1876 : 219, pi. 18, fig. 2 (as sp. of Heliconius). 1 6*. 1 9 syntypes. Godman- 
Salvin Coll. 1913-2. 1 q\ Santa Cruz, Peru. E. Bartlett. 1 $, Cosnipata Vail., E. Peru. 
H. Whitely. [PI. 6, fig. 67.] 

Currently placed as a subspecies of Heliconius aoede (Hubner) . 

batesi Riffarth, 1900 : 207 (as sp. of Heliconius). 6 $ syntypes. Original. / Ex. Coll. Riffarth. 
/ Joicey Bequest. Brit. Mus. 1934-120. 2 6*. Chanchamayo, Siid. Peru. 1 $, Peru? 3 o*> 
Ecuador. [PL 18, fig. 186.] 

Currently placed as a synonym of Heliconius melpomene xenoclea Hewitson. 

beata Riffarth, 19076 : 512, pi. 5, fig. 14 (as form of Heliconius erato eslrella). Holotype $. 
Ob. Pastaza, Ecuad., c. 1000 m. Cll. Niepelt. / Original. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 30, fig. 304.] 



TYPE-SPECIMENS OF HELICONIINAE 181 

Currently placed as a transitional form between Heliconius erato notabilis Salvin & Godman 
and Heliconius erato lativitta Butler. 

beebei Turner, 19666 : 128 (as subsp. of Heliconius demeter). Holotype <J, 2 $ paratypes. 
1 o\ J ?> British Guiana, Essiquibo R., 1 st. Falls. 14.x. 1929. Oxf. Univ. Expedn. B. M. 
1929-485. 1 $, Puruni River, B. Guiana. C. Buckle. / Ex. coll. C. J. Grist. 1925. [PI. 36, 
fig- 37 1 -] 

bella Riffarth, 19076 : 511 (as form of Heliconius cyrbia cyrbia). 1 $, 1 $ syntypes. Slanos, 
Ecuador. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, 
fig. 298.] 

Currently placed as a form of Heliconius erato cyrbia Godart. 

bicoloratus Butler, 1873a : 167 (as sp. of Heliconius). 2 <J syntypes. Nauta, Amazons 
Degand. 58-77. [PI. 10, fig. 108.] 

Currently placed as a form of Heliconius numata aristiona Hewitson. 

biedermanni Niepelt, 19266 : 337 (as form of Heliconius penelope). 3 c?. 1 ? syntypes. Joicey 
Bequest. Brit. Mus. 1934-120. 2 <J, i $, Bolivia. 1925. W. Niepelt, Zirlau. 1 <$, O. Bolivia. 
[PI. 25, fig. 254.] 

Currently placed as a transitional form between Heliconius melpomene amandus Grose- 
Smith & Kirby and Heliconius melpomene penelope Staudinger. 

broncus Stichel, 1906a : 21, pi. 2, fig. 6a (as subsp. of Heliconius cydno). 1 $ syntype. Peru? / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [Pi. 28, fig. 280.] 

Currently placed as a transitional form between Heliconius cydno alithea Hewitson and 
Heliconius cydno zelinde Butler. 

brunnea Stichel, 1903 : 6, pi. 1, fig. 5 (as ab. of Eueides isabella hippolina). 1 $ syntype. 
Tarapota, Peru. / Original. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 5, fig. 55.] 
Currently placed as a subspecies of Eueides isabella (Cramer). 

burchelli Poulton, 1910 : 331 (as subsp. of Heliconius nanna). 1 $ syntype, Chapada, Matto 
Grosso. H. H. Smith. / Godman-Salvin Coll. 1913-2. [PL 25, fig. 256.] 
Currently placed as a subspecies of Heliconius melpomene (L.). 

butleri Brown, 1975 : 217 (as subsp. of Heliconius paradalinus). Holotype o\ 4 6\ 3 ? paratypes. 
$ holotype, 1 $ paratype, Amazones, Perou, Cavallo-Cocho, Mai-Juillet 1884. M. de Mathan. / 
Levick Bequest. 1941-83. 1 <$, 1 $ paratype, Pebas, Amazones, M. de Mathan. fin Xbre & 
ier. Tr. 1880. / Levick Bequest. 1941-83. 1 $ paratype, Chambireyacu pres Yurimaguas, 
(Huallaga-Perou) M. de Mathan. Juin-Aout 1885. / Levick Bequest. 1941-83. 1 q" paratype, 
Iquitos, Peru, August 1931. (G. Klug). / Rothschild Bequest. B. M. 1939-1. 1 $ paratype, 
Upper Amazons, Iquitos, August 1932 (G. Klug). / RothschiM Bequest. B. M. 1 939-1. 1 $ 
paratype, Rio Pacaya, Lower Ucayali Peru. Aug. -Sept. 1912. / Joicey Bequest. Brit. Mus. 
I934" 120 - [Pi- J 3. ng. 136.] 

butleri Stichel, 1907 : 19 (as subsp. of Dione moneta). 3 $ syntypes. Typus. 2 q\ Colombia, 
R. Magdalena. e. c. H. Stichel. 1 o*. Ecuador, Balzapamba. e. c. H. Stichel. [PL 1, fig. 9.] 
Currently placed as a subspecies of Dione moneta Hiibner. 

caeruleatus Stichel, 1906a : 35 (as subsp. of Heliconius doris). 2 q* syntypes. Stid-Peru 
Pozuzo. e. c. H. Stichel. / Typus. 1 o* syntype, Rothschild Bequest. B. M. 1939-1. [PL 9, 
fig. 98.] 

Currently placed as a synonym of Heliconius doris doris (L.). 



182 P. R. ACKERY & R. L. SMILES 

calathus Stichel, 1909 : 178, fig. 2 (as subsp. of Eueides tales). 1 3" syntype. Ob. Pastaza 
Ecuad. c. 1000 m. Cll. Niepelt. Okt-Dec. 1906. / Cotype. / Joicey Bequest. Brit. Mus. 1934- 
120. [PI. 6, fig. 62.] 

Currently placed as a subspecies of Eueides tales (Cramer). 

callista Riffarth, 1900 : 188, 212 (as ab. of Heliconius phyllis). 1 $ syntype. Amazon super. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 31, fig. 320.] 

Currently placed as a synonym of Heliconius erato hydara form cally co-pis (Cramer). 

callycopis Cramer, [1777] : 143, pi. 190, figs E, F (as sp. of Papilio). 1 q" syntype. No. 20. 
Callicopis. Cr., II, 190, E. F. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 39, 

fig- 394-] 

Currently placed as a transitional form between Heliconius erato hydara Hewitson and 
Heliconius erato amalfreda Riffarth. 

candidus Brown, 1975 : 223 (as subsp. of Heliconius sapho). Holotype $, 5 6, 4 ? paratypes. 
o* holotype, Ecuador. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <J 
paratype, Ecuador, Fruhstorfer./ Joicey Bequest. Brit. Mus. 1934-120. 2 <§, 1 $ paratype, 
Ecuador. Hewitson Coll. 79.69. 1 $ paratype, Ecuador. / Ex. Grose Smith 1910 / Joicey. 
Bequest Brit. Mus. 1934-120. 1 $ paratype, Angamarca, Ecuador. Crowley Bequest. 1901-78. 
1 o* paratype, S. Domingo de los Colorados. W. Ecuad., October 1893 (W. Goodfellow). / 
Rothschild Bequest. B. M. 1939-1. 1 <$, 1 ? paratypes, Cachabe. low c. I. 97. (Rosenberg). 
Rothschild Bequest. B. M. 1939-1. [PI. 37, fig. 383.] 

carbo Stichel, 1903 : 13, pi. 1, fig. n (as form of Eueides lampelo acacetes). 1 o*. 1 $ 
syntypes. Santa Jnez (Ecuad.) . R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [P. 4, fig. 36.] 

Currently placed as a subspecies of Eueides lampeto Bates. 

carnea Neustetter, 1926a : 38 (as form of Heliconius penelope). Holotype $. Bolivia. 1925. 
W. Niepelt, Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 24, fig. 248.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 

carteri Riley, 1926 : 240, pi. 2, fig. 1 (as subsp. of Colaenis julia). 9 <J, 5 $ syntypes. 7$, 
Nassau, Bahamas. F. E. Taylor. / Godman-Salvin Coll. 1915-3. 1 o\ 1 $, Nassau, Bahamas. 
98-226. 16.10.98. 1 <J, 1 $, Bahamas. 1903-286. Sir G. Carter. 1 $, N. Providence, Bahama 
Is. F. E. Taylor. / Godman-Salvin Coll. 1915-3. 1 $, Bahamas, Nassau I. 7.x. 98. J. L. 
Bonhote. / Brit. Mus. 1924-188. 1 $, Bahamas. Sir G. Carter. 1904-200. [PL 2, fig. 21.] 
Currently placed as a subspecies of Dry as iulia (Fabricius). 

cassandra Felder & Felder, 18626 : 419 (as sp. of Heliconius). 1 $ syntype. Bogota. Lindig. 
Type. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PL 7, fig. 74.] 
Currently placed as a subspecies of Heliconius hecuba Hewitson. 

caternaulti Oberthiir, 1902 : 24, pi. 11, fig. 131 (as sp. of Heliconia). 3 6\ J ? syntypes. 
Levick Bequest. 1941-83. 2 (J, Guyane Francaise. Collection C. Bar. 1 q", Cayenne. 1 $, 
Bragance, (Para). M. de Mathan. [PL 7, fig. 80.] 

Currently placed as a synonym of Heliconius xanthocles vala Staudinger. 

catella Stichel, 1907 : 18, pi. 2, fig. 3 (as subsp. of Dione vanillae). Holotype $. Siid-Peru, 
Pozuzo. e. c. H. Stichel. / Rothschild Bequest. B. M. 1939-1. [PL 2, fig. 13.] 

Currently placed as a transitional form between Agraulis vanillae lucina Felder & Felder 
and Agraulis vanillae maculosa (Stichel). 

caucana Riley, 1926 : 242 (as subsp. of Colaenis euchroia). Lectotype $, 4 $ paralectotypes . 
Godman-Salvin Coll. 1915-3. <J lectotype, 3 (J paralectotypes, Frontino, Antioquia. T. K. 
Salmon. 1 $ paralectotype, Interior of Colombia, Wheeler. [PL 2, fig. 15.] 



TYPE-SPECIMENS OF HELICONIINAE 183 

Currently placed as a subspecies of Podotricha euchroia (Doubleday). 
Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 615. 

cephallenia Felder & Felder, 1865 : 373 (as sp. of Heliconius). 1 o* syntype. Surinam, 
Guiana. Beske. / DruceColl. Ex. KadenColl. / Godman-Salvin Coll. 1913-2. [PI. 17, fig. 170.] 
Currently placed as a subspecies of Heliconius ethilla Godart. 

ceres Oberthur, 1920 : 30, pi. 519, fig. 4324 (as sp. of Heliconia). 2 0" syntypes. Nouvelle 
Granade, Cauca, Juntas. M. de Mathan. fin 1897, 1-1898. / Levick Bequest. 1941-83. [PL 38, 
fig. 388.] 

Currently placed as a transitional form between Heliconius eleuchia eleuchia Hewitson and 
Heliconius eleuchia eleusinus Staudinger. 

chapadensis Brown, 1973 : 6 (as subsp. of Heliconius ethilla). 3 q*, 2 $ paratypes. 1 o* 
paratype, Matto Grosso, Brazil. / Brit. Mus. 1933-161. 1 o* paratype, Amazons. H. H. 
Smith. / Godman-Salvin Coll. 1913-2. 1 $, 1 $ paratypes, Burity, 30 miles N. E. [48 km] 
of Cuyaba 2250 ft [686 m] 17-30. vi. 27. Matto Grosso. C. L. Collenette / Joicey Bequest. 
Brit. Mus. 1934-120. 1 $ paratype, Burity, 30 miles N. E. [48 km] of Cuyaba 2250 ft [686 km] 
1-14.vii.27. Matto Grosso. C. L. Collenette / Joicey Bequest. Brit. Mus. 1934-120. [PI. 17, 
fig. 171.] 

chestertonii Hewitson, 18726 : fig. 22 (as sp. of Heliconia). 1 <$, 2 $ syntypes. N. Granada. 
Hewitson Coll. 79. 69. [PI. 29, fig. 294.] 

Currently placed as a subspecies of Heliconius erato (L.). 

chioneus Bates, 1864a : 58 (as sp. of Heliconius). Holotype o". Lion Hill, Panama, 
McLeannan. / Godman-Salvin Coll. 1913-2. [PI. 26, fig. 263.] 
Currently placed as a subspecies of Heliconius cydno Doubleday. 

choarina Hewitson, 1872a : 83 (as sp. of Heliconia). 1 $ syntype. Ecuador, Hewitson 
Coll. 79.69. [PI. 7. fi g- 77-J 

Currently placed as a subspecies of Heliconius hecuba Hewitson. 

chrysantis Godman & Salvin, 1881 : 146 (as sp. of Heliconius). Holotype o"- Nicaragua, 
T. Bridges. / Godman-Salvin Coll. 1913-2. [PL 14, fig. 141.] 

Currently placed as a transitional form between Heliconius hecale zuleika Hewitson and 
Heliconius hecale fornarina Hewitson. 

clarescens Butler, 1875 : 223 (as sp. of Heliconius). Holotype $. Bugaba, Panama. Arce. / 
Godman-Salvin Coll. 191 3-2. [PL 12, fig. 131.] 

Currently placed as a subspecies of Heliconius ismenius Latreille. 

claudia Godman & Salvin, 1881 : 145 (as sp. of Heliconius). 2 o* syntypes. Calobre, Panama, 
Arce. / Godman-Salvin Coll. 191 3-2. [PL 16, fig. 168.] 

Currently placed as a subspecies of Heliconius ethilla Godart. 

clearei Hall, 1930 : 278 (as subsp. of Heliconius hecale). 10 <$, 4 $ syntypes. Mabaruma, 
British Guiana, Dec. 1929-Jan. 1930. A. Hall. 4 <J, 2 $, Joicey Bequest. Brit. Mus. 1934-120. 
5 o", 2 $, Brit. Mus. 1930-567. [PL 16, fig. 161.] 

clearista Oberthur, 1923 : 304, pi. 565, fig. 4863 (as ab. of Heliconius egeria). 1 q* syntype. 
Guyane francaise. / Levick Bequest. B. M. 1941-83. [PL 8, fig. 91.] 
Currently placed as an aberration of Heliconius egeria egeria (Cramer) . 

clelia Neustetter, 1927a : 86 (as form of Heliconius phyllis anacreon). Holotype o*- Bolivia. / 
Joicey Bequest. Brit. Mus. 1934-120. [PL 34, fig. 345.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius) . 



1 84 P. R. ACKERY & R. L. SMILES 

clytie Neustetter, ig2jb : 229 (as form of Heliconius xenoclea plesseni). Holotype q*- 
Guayes, Ecuador. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 20, fig. 199.] 

Currently placed as a transitional form between Heliconius melpomene plesseni Riffarth 
and Heliconius melpomene aglaope Felder & Felder. 

cognata Riffarth, 19076 : 506 (as form of Heliconius melpomene aglaope). 4 $ syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 o*. Pozuzo, Peru. 
800 m. Hoffman. 1 o*. Peru. H. Fruhstorfer. 2 $, Peru, Pozuzo. H. Fruhstorfer. [PI. 20, 
fig. 204.] 

Currently placed as a form of Heliconius melpomene aglaope Felder & Felder. 

collis Joicey & Kaye, 1917a : 413, pi. 57, fig. 3 (as ab. of Heliconius melpomene melpomene). 
Holotype q\ Guyane Francse., Nouveau Chantier. Collection Le Moult. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 2i, fig. 215.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

compact a Joicey & Kaye, 1919 : 349 (as ab. of Heliconius melpomene melpomene). 
Holotype $. Guyane Francse., St. Jean du Maroni. Collection Le Moult. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 21, fig. 216.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene meriana Turner. 

completa Oberthiir, 1920 : 37, pi. 522, fig. 4334 (as form of Heliconia cethosia). 2 <J syntypes. 
1 o\ Nouvelle-Grenade, Region de Bogota. Recu en decembre 191 7 du frere Apollinaire-Marie. / 
Joicey Bequest. Brit. Mus. 1934-120. 1 o*> Nouvelle-Grenade, Env. de Bogota. Frere 
Apollinaire-Marie. recu 2 avril 1917. / Levick Bequest. 1941-83. [PI. 8, fig. 82.] 
Probably best regarded as a synonym of Heliconius xanthocles flavosia Kaye. 

concinna Stichel, 1906a : 28 (as form of Heliconius vulcanus). Holotype o*- Balzapamba 
(Ecuad.). R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934- 
120. [PI. 18, fig. 184.] 

Currently placed as a form of Heliconius melpomene cythera Hewitson. 

confluens Lathy, 1906 : 453, pi. 34, fig. 3 (as var. of Heliconius xenoclea). 1 £, 1 $ syntypes. 
Adams Bequest B. M. 1912-399. 1 $, Pichis Road, Peru. 3000 ft [914 m] 04. Watkins and 
Tomlinson. 1 ?, Rio Colorado, Peru, 2500 ft [762 m] 04. Watkins and Tomlinson. [PL 18, 
fig. 185.] 

Currently placed as a transitional form between Heliconius melpomene xenoclea Hewitson 
and Heliconius melpomene euryades Riffarth. 

confluens Neustetter, 19280" : 259 (as form of Heliconius cydno cydnides). 1 $ syntype. 
Solano, Yapura. / Joicey Bequest. Brit. Mus. 1934-120. [PL 26, fig. 266.] 

Currently placed as a transitional form between Heliconius cydno zelinde Butler and 
Heliconius cydno cydnides Staudinger. 

constricta Joicey & Kaye, 1917a : 427 (as ab. of Heliconius erato). Holotype <J. Guyane 
Francse., St. Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 32, fig. 329.] 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara Hewitson. 

copiosus Stichel, 1906a : 57 (as subsp. of Eueides lampeto). 2 $ syntypes. Potaro Road, 
Potaro Essequibo, Brit. Guiana. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. [PL 3, fig. 31.] 

coralii Butler, 1877 : 151 (as sp. of Heliconius). 2 o\ 2 9 syntypes. Amazons Trail. 77-64. 
1 o\ Serpa Bush 25.4.74. z 6*. l ?> SerpaBush, 21.4.74. * ?< Serpa, 13.75. [PL 32, fig. 324.] 



TYPE-SPECIMENS OF HELICONIINAE 185 

Currently placed as a transitional form between Heliconius erato amalfreda Riffarth and 
Heliconius erato hydara Hewitson. 

corona Niepelt, 1907a 1213 (as form of Heliconius batesi plesseni). 1 o* syntype. Original. / 
Adams Bequest. B. M. 1912-399. [PI. 19, fig. 195.] 

Currently placed as a form of Heliconius melpomene plesseni Riffarth. 

crespinus Kriiger, E. G., 1925 : 151 (as var. of Heliconius crispus). 1 $ syntype. 24.6.16. 
2200. Cauca V. / Cotype. / Collection Niepelt. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 7, fig. 76.] 

Currently placed as a form of Heliconius hecuba crispus Staudinger. 

crystalina Hall, 192 1 : 279 (as sp. of Eueides). 7 £, 2 $ syntypes. Crystalina, W. Colombia. 
1 100 ft, [330 m] June-July 1920. A. Hall. 3 o\ Brit. Mus. 1929-292. 4 <$, 2 2, Joicey 
Bequest. Brit. Mus. 1934-120. [PI. 6, fig. 61.] 

Currently placed as a synonym of Eueides tales xenophanes Felder. 

cupidineus Stichel, 1906a : 31 (as subsp. of Heliconius aeode). 1^,1$ syntypes. Original / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 $, Tarapoto, Peru. 1 $, 
Juanjuy, Peru. [PI. 6, fig. 68.] 

curvifascia Talbot, 1928 : 220 (as form of Heliconius melpomene burchelli). Holotype <J. 
Melguira, 10 miles S. [16 km] of Diamantino, 2000 ft [608 m] 23.v-.-3.vi.27. Matto Grosso. 
C. L. Collenette. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 257.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene burchelli Poulton. 

cybeleia Joicey & Kaye, 1919 : 349 (as ab. of Heliconius melpomene cybele). Holotype $. 
Guyane Francse., St-Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 22, fig. 221.] 

Probably best regarded as a synonym of Heliconius melpomene meriana form obscurata 
Riffarth. 

cybelellus Joicey & Kaye, 191 76 : 92, pi. 6, fig. 4 (as ab. of Heliconius erato erato). 1 q* syntype. 
Serpa, Lower Amazon, Jan-March, 1914. A. Hall. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 32, fig. 325.] 

Currently placed as a transitional form between Heliconius erato amalfreda Riffarth and 
Heliconius erato hydara Hewitson. 

cydno Doubleday, 1847 : 103, pi. 15, fig. 3 (as sp. of Heliconia). 1 ? syntype. Bogota, Miers. 
46-55. [PI. 26, fig. 264.] 

Currently placed as a species of Heliconius Kluk. 

cythera Hewitson, 18696 : 9 (as sp. of Heliconia). 2 $ syntypes. Ecuador. Hewitson Coll. 
79.69. [PI. 18, fig. 183.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 

deflava Joicey & Kaye, 19176 : 93, pi. 6, fig. 6 (as ab. of Heliconius sapho primularis). 2 <$, 
1 ? syntypes. Paramba, Ecuador. Rosenburg. / Ex Grose Smith, 1910. / Joicey Bequest. 
Brit. Mus. 1934-120. (PL 38, fig. 386.] 

Currently placed as a transitional form between Heliconius eleuchia primularis Butler and 
Heliconius eleuchia eleusinus Staudinger. 

deleta Stichel, 1907 : 13 (as form of Colaenis phaetusa). Holotype q*. Paraguay, Nuev. 
Germania, e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M 1939-1. [PI. 1, fig. 4.] 
Currently placed as a form of Dryadula phaetusa (L.). 

diana Riffarth, 1900 : 204 (as ab. of Heliconius melpomene). Holotype $. Cayenne? / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 229.] 



186 P. R. ACKERY & R. L. SMILES 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene meriana Turner. 

dianides Joicey & Kaye, 1917a : 417 (as ab. of Heliconius melpomene cybele). Holotype o*- 
Guyane Francse, St Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934- 
120. [PI. 22, fig. 225.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene meriana Turner. 

diaphana Niepelt, 1915 : 58 (as var. of Colaenis tithraustes). Holotype q\ S. O. Peru. 1903. / 
Collection Niepelt. / Presented by J. J. Joicey Esq. Brit. Mus. 1931-291. [PI. 2, fig. 20.] 
Currently placed as a synonym of Podotricha telesiphe telesiphe (Hewitson). 

diatonica Fruhstorfer, 1912 : 14 (as subsp. of Metamandana dido). Holotype o*. Honduras, 
San Pedro Sula. ex. coll. Fruhstorfer. / Fruhstorfer Coll. B. M. 1937-285. [PI. 1, fig. 2.] 

Currently placed as a synonym of Philaethria dido (L.), but may eventually be recognized 
as a good species. 

diffluens Riffarth, 19076 : 513 (as form of Heliconius erato phyllis). 3 o*. 4 ? syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 2 0". 2 ?, Prov. Sara, 
Dep. St Cruz, Siid Bolivia. 1 q", i $, Peru. 1 $, Slid Brazilien. [PI. 34, fig. 348.I 

diffusus Butler, 1873a : 168 (as sp. of Heliconius). 1 o* syntype. Para, Bates 50-4. [PI. 12, 
fig. 128.] 

Currently placed as a form of Heliconius numata silvana (Cramer). 

diformata Riffarth, 1900 : 209 (as ab. of Heliconius cyrbia). 1 o* syntype. Ecuador, Quito. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, fig. 297.] 

Currently placed as a transitional form between Heliconius erato cyrbia Godart and 
Heliconius erato venus Staudinger. 

diotrephes Hewitson, 1869a : 33 (as sp. of Heliconia). 2 oV 2 $ syntypes. Nicaragua. 
Hewitson Coll. 79.69. [PI. 26, fig. 261.] 

Probably best regarded as a synonym of Heliconius cydno galanthus Bates. 

dissoluta Stichel : 1903 : 6, pi. 1, fig. 7 (as subsp. of Eueides isabella). 1 <$, 1 $ syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 o\ Tarapota, Peru. 
1 $, Bolivien. [PI. 5, fig. 56.] 
Currently placed as a subspecies of Eueides isabella (Cramer). 

diva Stichel, 1906a : 47 (as subsp. of Heliconius erato). Holotype $. Bolivia. / Original. / 

Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 33, fig. 340.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato lativitta Butler. 

dryope Riffarth, 1900 : 212 (as ab. of Heliconius phyllis). 1 o* syntype. Surinam, H. Stichel. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. /Brit. Mus. 1934-120. [PI. 32, fig. 323.] 

Currently placed as a transitional form between Heliconius erato amalfreda Riffarth and 
Heliconius erato hydara Hewitson. 

dynastes Felder & Felder, 1861 : 102 (as sp. of Eueides). 1 o", 1 $ syntypes. Felder Colin. / 
Rothschild Bequest. B. M. 1 939-1. 1 6*. Bogota. Lindig. Type. 1 $, Venezuela. Moritz. 
Type. [PL 5, fig. 57.] 

Currently placed as a form of Eueides isabella huebneri Menetries. 

eanes Hewitson, 1861 : 155, pi. 10, fig. 1 (as sp. of Eueides). i-<J syntype. Amazons, Hewitson 
Coll. 79-69. [PL 4. fi g- 45-] 



TYPE-SPECIMENS OF HELICONIINAE 187 

eanides Stichel, 1903 : 30, pi. 1, fig. 24 (as ab. of Eueides eanes). 1 6* syntype. Bolivien. / 
Original. / Ex. Coll. Riffarth. [PI. 5, fig. 47.] 

Currently placed as a synonym of Eueides eanes heliconioides Felder & Felder. 

edias Hewitson, 1861 : 155, pi. 10, fig. 2 (as sp. of Eueides). 4 q\ i $ syntypes. New Grenada. 
Hewitson Coll. 79. 69. [PI. 3, fig. 26.] 

Currently placed as a subspecies of Eueides procula Doubleday. 

egeriformis Joicey & Kaye, 1917a : 430, pi. 107, fig. 8 (as sp. of Eueides). 5 q* syntypes. 
Joicey Bequest. Brit. Mus. 1934-120. 4 q\ Guyane Francse, St Jean du Maroni. Collection 
Le Moult. 1 o\ Guyane Francse. Nouveau Chantier. Collection Le Moult. [PI. 36, fig. 369. J 
Currently placed as a synonym of Heliconius demeter bouqueti Noldner. 

cgregia Riffarth, 19076 : 505 (as form of Heliconius cydno alithea). 2 <J syntypes. Balzapamba 
(Ecuad.). R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934- 
120. [PI. 28, fig. 284.] 

elegantula Joicey & Kaye, 1917a : 417 (as ab. of Heliconius melpomene cybele). Holotype (J. 
Guayane Fr., St. Jean, vii-ix.15. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 22, fig. 227.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

eleuchia Hewitson, [1853] : pi. 3, fig. 8 (as sp. of Heliconia). 1 <J, i $ syntypes. N. Granada. 
Hewitson Coll. 79.69. [PI. 37, fig. 385.] 

Currently placed as a species of Heliconius Kluk. 

elsa Riffarth, 1899 : 407 (as var. of Heliconius clytia). 1 <J syntype. Surinam, ex. coll. 
Fruhstorfer. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 8, 

fig- 85.] 

Currently placed as a form of Heliconius wallacei wallacei Reakirt. 

eltringhami Joicey & Kaye, 1917a : 418, pi. 107, fig. 6 (as subsp. of Heliconius melpomene). 
Holotype q*- St. Laurent, Maroni River, French Guiana. July-Sept. 1915. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 2i, fig. 211.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

emilius Weymer, 1912 : 73 (as sp. of Heliconius). Holotype $. Muzo, Colombia 400 b. 
800 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 27, fig. 278.] 

Currently placed as a hybrid between Heliconius melpomene melpomene (L.) and Heliconius 
cydno cydno Doubleday. 

etnma Riffarth, 1901 : 164 (as subsp. of Heliconius phyllis). 1 <J, 3 ? syntypes. Original. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <J, Amazon super. 2 $, Yurim, 
Peru. Mich. 1 $, Archidona (Ecuad.). R. Haensch S. [PI. 30, fig. 309.] 
Currently placed as a subspecies of Heliconius evato (L.). 

emmelina Oberthiir, 1902 : 24, pi. 11, fig. 132 (as sp. of Heliconia). Holotype (J. Guyane 
anglaise. / Levick Bequest. B. M. 1941-83. [PI. 6, fig. 66.] 

Currently placed as a synonym of Heliconius aoede astydamia Erichson. 

emsleyi Brown, 1975 : 211 (as sp. of Eueides). Holotype <$, 1 ? paratype. 1 q\ Valdevia 
Colombia. Pratt, 97. / Ex. Grose Smith, 1910. 1 $, Equateur, La Chima. M. de Mathan 
1 Semestre 1893. / Ex. Oberthiir Coll. Brit. Mus. 1927-3. [PI. 4, fig. 38.] 

eratophylla Joicey & Kaye, igiyb : 91, pi. 6, fig. 2 (as subsp. of Heliconius evato). Holotype $. 
Tarapo, Peru, S. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, fig. 299.] 

Currently placed as a transitional form between Heliconius evato favovinus Hopffer and 
Heliconius evato lativitta Butler. 



188 P. R. ACKERY & R. L. SMILES 

eratosignis Joicey & Talbot, 1925 : 649 (as sp. of Eueides). 1 <J, 1 $ syntypes. Joicey Bequest. 
Brit. Mus. 1934-120. 1 $, River System Cuyaba-Corumba, Matto Grosso, Brazil. [PI. 36, 

fig- 372] 

Currently placed as a subspecies of Heliconius demeter Staudinger. 

erythrea Cramer, [1777] : 140, pi. 189, fig. A (as sp. of Papilio). 1 <J syntype. No. 14. 
Erythrea. Cr., II, 189, A. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PL 39, 

fig- 395-1 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara Hewitson. 

estrella Bates, 1862 : 560 (as sp. of Heliconius). 1 <$, 1 $ syntypes. Godman-Salvin Coll. 
1913-2. 1 <J, Marajo I., Amazons. H. W. Bates. 1 $, Para, L. Amazons. H. W. Bates 
[PI. 33, fig. 334.] 

Currently placed as a subspecies of Heliconius erato (L.). 

etylus Salvin, 1871 : 414 (as sp. of Heliconius). 1 <$ syntype. Gualaquiza, Ecuador. Pearce. / 
Godman-Salvin Coll. 191 3-2. [PI. 30, fig. 308.] 

Currently placed as a transitional form between Heliconius erato lativitta Butler and 
Heliconius erato notabilis Salvin & Godman. 

eucherius Weymer, 1906 : 68 (as sp. of Heliconius). Holotype <$. Bogota, Columbien. / Ex. 
Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 14, fig. 149.] 
Currently placed as a form of Heliconius hecale holcophorus Staudinger. 

euchroia Doubleday, 1847 : pi. 20, fig. 3 (as sp. of Colaenis). Lectotype <£. Columbia. 
Bought from Jurgenson. 43-64. [PI. 2, fig. 14.] 

Currently placed as a species of Podotricha Michener. 

Described from Venezuela, New Granada. 

Lectotype designated by Vane- Wright, Ackery & Smiles, 1975 : 613. 

eueidina Oberthiir, 1916 : 37, pi. 406, fig. 3487 (as sp. of Heliconia). 2 £ syntypes. Levick 
Bequest. 1941-83. 1 $, Guyane Francaise. Collection C. Bar. 1 $, Ex. Musaeo Boisduval. / 
Cayenne. [PI. 36, fig. 370.] 

Currently placed as a synonym of Heliconius demeter bouqueti Noldner. 

euphone Felder & Felder, 18626 : 418 (as sp. of Heliconius). 2 $ paralectotypes. Bogota. 
Lindig. / Felder Colin. / Rothschild Bequest. 1939-1. [PI. 10, fig. 102.] 
Currently placed as a subspecies of Heliconius numata (Cramer). 
Lectotype designated by Holzinger & Holzinger, 1974 : 264. 

euryades Riffarth, 1900 : 205 (as var. of Heliconius amaryllis). 1 <J syntype. Vilcanota, 
3000 ft [914 m] Peru. Garlepp. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
\q^-\zo. [PL 24, fig. 245.] 
Currently placed as a subspecies of Heliconius melpomene (L.). 

eurycleia Brown, 1973 : 4 (as subsp. of Heliconius aoede). 1 $, 1 $ paratypes. Areia Branca, 
MT. Km. 575, Cuiaba-Porto Velho. 23.vi.1971. / Brit. Mus. 1974-63. [PL 6, fig. 69.] 

eurysaces Hewitson, 1864 : 248, pi. 16, fig. 3 (as sp. of Eueides). 1 $ syntype. Quito. 
Hewitson Coll. 79.69. [PL 3, fig. 29.] 

Currently placed as a subspecies of Eueides procula Doubleday. 

excellens Neustetter, 1926a : 37 (as form of Heliconius penelope). Holotype $. Bolivia. 1925. 
W. Niepelt. / Joicey Bequest. Brit. Mus. 1934-120. [PL 24, fig. 247.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 



TYPE-SPECIMENS OF HELICONIINAE 189 

exornata Riffarth, 19076 : 505 (as form of Heliconius cydno galanthus). 4 q* syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 2 <J, Limon, Costa 
Rica. Sept. -Oct. 1904. A. Hall. 1 (J, Carillo, Costa Rica. 1 o*. San Carlos, Costa Rica. 
[PI. 26, fig. 262.] 

Currently placed as a transitional form between Heliconius cydno galanthus Bates and 
Heliconius cydno chioneus Bates. 

faivrei Joicey & Kaye, 1919 : 349 (as ab. of Heliconius melpomene melpomene). Holotype $. 
Guyane Francse., St. -Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PL 21, fig. 218.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meviana Turner. 

fasciatus Godman & Salvin, 1877 : 62 (as sp. of Heliconius). 3 <J syntypes. Lion Hill, 
Panama. McLeannan. / Godman-Salvin Coll. 1913-2. [PL 13, fig. 135.] 
Currently placed as a form of Heliconius ismenius ismenius Latreille. 

faustalia Joicey & Kaye, 1917a : 417 (as ab. of Heliconius melpomene cybele). Holotype $. 
Guyane Francse., St-Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934- 
120. [PL 22, fig. 223.] 

Currently placed as a transitional form of Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meviana Turner. 

fischeri Fassl, 191 2 : 55 (as subsp. of Heliconius clysonimus). Holotype $. Rio Aguacatal, 
Columb., W. Codr. 2000 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. [PL 36, 

fig- 364] 

Currently placed as a transitional form between Heliconius clysonymus clysonymus 
Latreille and Heliconius clysonymus hygiana Hewitson. 

flagrans Stichel, 1919 : 120 (as subsp. of Heliconius amaryllis). 2 q* syntypes. Trinidad, 
Port of Spain. / Typus. / e. c. H. Stichel. /Rothschild Bequest. B. M. 1939-1. [PL 21, 
fig. 210.] 

Currently placed as a form of Heliconius melpomene melpomene (L.). 

flammea Niepelt, 1925 : 252 (as form of Heliconius phyllis). Holotype o*- Bolivia. 1925. 
W. Niepelt, Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PL 25, fig. 253.] 

Currently placed as a transitional form between Heliconius melpomene amandus Grose- 
Smith and Heliconius melpomene penelope Staudinger. 

flava Neustetter, 1928c : 247 (as form of Heliconius hecuba choarinus). Holotype $. Baiza 
(Ecuad.). R. Haensch. 1500 m. 16. i. 00. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. [PL 7, fig. 78.] 

Currently placed as a form of Heliconius hecuba choarina Hewitson. 

Described from Ecuador, Macas. 

flaveola Joicey & Kaye, 19176 : 93, pi. 6, fig. 3 (as subsp. of Heliconius cydno). 1 £ syntype. 
Mocotone, Venezuela. / Ex. Coll. Herbert Druce, 1913 / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 38, fig. 389.] 

Currently placed as a transitional form between Heliconius cydno perijaensis Masters and 
Heliconius cydno barinasensis Masters. 

flavidior Neustetter, 1928^ : 258 (as form of Heliconius cydno cydnides). Holotype <$. 
Solano, Yapura. / Joicey Bequest. Brit. Mus. 1934-120. [PL 27, fig. 271.] 
Probably best regarded as a synonym of Heliconius cydno zelinde Butler. 

flavopunctatus Fassl, 1912 : 55 (as subsp. of Heliconius clysonimus). 1 o*> * ? syntypes. Rio 
Aguacatal, Columb., W. Codr., 2000 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 35, fig. 362.] 



igo P. R. ACKERY & R. L. SMILES 

Currently placed as a transitional form between Heliconius clysonymus clysonymus Latreille 
and Heliconius clysonymus hygiana Hewitson. 

flavorubra Neustetter, 19266 : 279 (as form of Heliconius melpomene) 1 <J syntype. St. 
Laurent, Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 21, fig. 213.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene meriana Turner. 

fortnosus Bates, 1866 : 87 (as sp. of Heliconius). 1 $ syntype. Lion Hill, Panama. 
McLeannan. / Godman-Salvin Coll. 1913-2. [PI. 35, fig. 355.] 

Currently placed as a subspecies of Heliconius hecalesia Hewitson. 

fornarina Hewitson, [1853] : pi. 3, fig. 9 (as sp. of Heliconia). 1 <J syntype. S. America. 
Capt. Belcher. 44-6. [PI. 13, fig. 139.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius). 

framptoni Riley, 1926 : 241, pi. 2, fig. 5 (as subsp. of Colaenis julia). 3 <J, 2 $ syntypes. 2 o", 
West Indies, St. Vincent. April, 1925. R. E. E. Frampton. / Brit. Mus. 1925-529. 1 <J, 
1 $, Windward side, St. Vincent, W. I., H. H. Smith. / Godman-Salvin Coll. 1915-3. 1 $, 
St. Vincent, W. I., H. H. Smith. / Godman-Salvin Coll. 1915-3. [PI. 3, fig. 23.] 
Currently placed as a subspecies of Dry as iulia (Fabricius) . 

fraterna Niepelt, 1914 : 40 (as form of Heliconius melpomene aglaope). 1 £ syntype. 
Ecuador, Canelos. 800 m. / Original. / Adams Bequest. B. M. 1912-399. [PI. 19, fig. 198.] 
Currently placed as a form of Heliconius melpomene plesseni Riffarth. 

fruhstorferi Riff arth, 1899 : 406 (as sp. of Heliconius). Holotype $. Espirito-Santo, Brasil. 
ex. coll. Fruhstorfer. / Original. / Ex. Coll. Riffarth. [PI. 9, fig. 100.] 
Currently placed as a synonym of Heliconius nattereri Felder & Felder. 

fulgidus Stichel, 1906a : 41 (as subsp. of Heliconius sard). 5 $, 1 $ syntypes. 1 <$, 1 $, Costa- 
Rica (Fr. Schneider), e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. 1 $, 
Costa-Rica, Carillo. e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. 1 $, 
Carillo, Costa Rica. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
1 o*. Costa Rica. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 $, 
Piedras Negras. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 37, fig. 375.] 

Probably best regarded as a synonym of Heliconius sara say a (Fabricius). 

fuliginosa Riffarth, 19076 : 511 (as form of Heliconius erato erato). 2^,4? syntypes. Original. 
/ Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <$, Surinam? 1 <J, 2 $, Berg 
en Daal, Sur., Juli. Michaelis. 2 $, Cayenne. [PI. 32, fig. 328.] 

fuliginosa Stichel, 1903 : 12, pi. 1, fig. 10 (as ab. of Eueides lampeto). 1 £, 1 $ syntypes. Ex. 
Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 £, Santa Jnez (Ecuad.). R. Haensch 
S. / Original. 1 $, Santa Jnez (Ecuad.). R. Haensch S.. 1 $, Bafios (Ecuad.). R. Haensch 
S. [PI. 3, ng. 33.] 

Currently placed as a form of Eueides lampeto acacetes Hewitson. 

fulvescens Lathy, 1906 : 452, pi. 34, fig. 1 (as var. of Heliconius pasitkoe). Holotype $. 
Demerara, Castell. / Adams Bequest. B. M. 1912-399. [PL 16, fig. 159.] 
Currently placed as a form of Heliconius hecale hecale (Fabricius). 

galanthus Bates, 1864a : 58 (as sp. of Heliconius). 3 6\ 2 $ syntypes. Godman-Salvin Coll. 
1913-2. 1 <J, Montagua Valley. F. D. G. & O. S. 1 6", 1 ?, Forests of N. Vera Paz. F. D. G. & 
O. S. 1 o\ 1 $, Choctum, Guatemala. F. D. G. & O. S. [PL 26, fig. 260.] 
Currently placed as a subspecies of Heliconius cydno Doubleday. 



TYPE-SPECIMENS OF HELICONIINAE 191 

gibbsi Kaye, 1919 : 217 (as ab. of Heliconius doris doris). Holotype <J. Friendship, B. Guiana. 
/ Ex. Coll. Schmassmann, 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 9, fig. 97.] 
Currently placed as a form of Heliconius doris doris (L.). 
Described from a single female specimen only; however, this male is apparently the type. 

glaucina Neustetter, 19286 : 80 (as form of Heliconius estrella). Holotype $. S. Columbien, 
Mocoa. November 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 31, fig. 312.] 

Currently placed as a transitional form between Heliconius erato dignus Stichel and 
Heliconius erato lativitta Butler. 

gnophota Stichel, 1907 : 20, pi. 2, fig. 4 (as subsp. of Dione glycera). 1 <J, 1 ? syntypes. 
Colombia, Cordill. p. Bogota, e. c. H. Stichel. / Rothschild Bequest. B. M. 1 939-1. [PI. 1, 
fig. 8.] 

Currently placed as a synonym of Dione glycera (Felder & Felder) . 

gracilis Riffarth, igoyb : 504, pi. 5, fig. 3 (as form of Heliconius aristiona lepidus). 1 <J, 1 $ 
syntypes. Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q\ La 
Merced, Peru. 2500 ft [762 m] '04. 1 $, Tarapoto, Peru. [PI. 10, fig. 104.] 
Currently placed as a form of Heliconius numata euphone Felder & Felder. 

gracilis Stichel, 1903 : 23 (as subsp. of Eueides aliphera). 2 q* syntypes. Typus. / Rothschild 
Bequest. B. M. 1939-1. 1 $, Honduras, San Pedro Sula e. c. H. Stichel. 1 ^, Costa-Rica 
San Carlos e. c. H. Stichel. [PI. 6, fig. 63.] 

guiensis Riffarth, 1900 : 198 (as ab. or var. of Heliconius numata). Holotype $. British 
Guayana. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 11, 
fig. 122.] 

Currently placed as a form of Heliconius numata numata (Cramer). 

gustavi Staudinger, 1896 : 287 (as ab. of Heliconius weymeri). 1 $ syntype. Rio Dagua, 
Columb. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 27, fig. 276.] 
Currently placed as a form of Heliconius cydno weymeri Staudinger. 

gynaesia Hewitson, 1875 : 182 (as sp. of Heliconia). Holotype (J. Hewitson Coll. 79.69. 
[PI. 35. ng. 356.] 

Currently placed as a subspecies of Heliconius hecalesia Hewitson. 

haenschi Riffarth, 1900 : 200 (as ab. of Heliconius cydno). 1 $, 1 $ syntypes. Balzapamba 
(Ecuad). R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934- 
120. [PI. 28, fig. 281.] 

Currently placed as a form of Heliconius cydno alithea Hewitson. 

halli Kaye, 1919 : 217 (as ab. of Heliconius wallacei brevimaculata) . Holotype $. Serpa, 
Lower Amazon. March 1914. A. Hall. / Ex. Coll. Schmassmann, 1927. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 8, fig. 88.] 

Currently placed as a form of Heliconius wallacei wallacei Reakirt. 

hecuba Hewitson, 1858 : pi. 4, fig. (as sp. of Heliconia). 4 $ syntypes. N. Granada, Hewitson 
Coll. 79.69. [PI. 7, fig. 72.] 

Currently placed as a species of Heliconius Kluk. 

Helena Riffarth, 19076 : 511 (as form of Heliconius erato cybelinus). 2 £ syntypes. Original. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <$, Obidos, Amazonen strom. 
viii.xi.99. ex. coll. Fruhstorfer. 1 £, Berg en Dal, Sur., Juli. Michaelis. [PI. 32, fig. 321.] 

Currently placed as a transitional form between Heliconius erato amalfreda Riffarth and 
Heliconius erato hydara Hewitson. 

heliconioides Felder & Felder, 1865 : 378, pi. 46, fig. 16 (as sp. of Eueides). 1 <J syntype. 
Ecuador, type. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 6, fig. 65.] 
Currently placed as a subspecies of Eueides eanes Hewitson. 



192 P. R. ACKERY & R. L. SMILES 

hemicycla Joicey & Kaye, 1917a : 427 (as ab. of Heliconius erato). 6 $ syntypes. Guyane 
Francse., St. Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 32, fig. 327.] 

Currently placed as a form of Heliconius erato erato (L.). 

henrici Kriiger, R., 1929 : 376 (as form of Heliconius phyllis anactorie). Holotype $. Carmen, 
Rio Madre de Dios, Nord Bolivia. / Levick Bequest. B. M. 1941-83. [PL 34, fig. 342.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius) . 

heraldicus Stichel, 1903 : 27, pi. 1, fig. 19 (as subsp. of Eueides thales). 1 $ syntype. Para. 
10.92. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 6, 
fig. 60.] 

Currently placed as a synonym of Eueides tales pythagoras Kirby. 

hermanni Riffarth, 1899 : 407 (as var. oder ab? of Heliconius ismenius). Holotype $. 
Columb. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 13, 

fig- 134] 

Currently placed as a form of Heliconius ismenius ismenius Latreille. 

hermathena Hewitson, [1853] : pi. 2, fig. 5 (as sp. of Heliconia). 1 $ syntype. Amazons. 
Hewitson Coll. 79.69. [PL 29, fig. 290.] 

Currently placed as a species of Heliconius Kluk. 

hermogenes Hewitson, 1858 : pi. 4, fig. 12 (as sp. of Heliconia). 1 $ syntype. Ex. Musaeo 
Boisduval. / Levick Bequest. 1941-83. [PL 27, fig. 269.] 

Currently placed as a subspecies of Heliconius cydno Doubleday. 

hierax Hewitson, 18696 : 11 (as sp. of Heliconia). 3 o*. * ? syntypes. Ecuador. Hewitson 
Coll. 79.69. [PL 7, fig. 71.] 

Currently placed as a species of Heliconius Kluk. 

himera Hewitson, 1867a : pi. 5, fig. 16 (as sp. of Heliconia) . 3 <$ syntypes. Ecuador. 
Hewitson Coll. 79.69. [PL 29, fig. 291.] 

Currently placed as a species of Heliconius Kluk. 

hippola Hewitson, 1867a : pi. 5, fig. 13 (as sp. of Heliconia) . 1 $ syntype. Hewitson Coll. 
79.69. [PL 17, fig. 173.] 

Currently placed as a hybrid between Heliconius melpomene melpomene (L.) and Heliconius 
ethilla metalilis Butler. 

hippolinus Butler, 1873a : 169 (as sp. of Eueides). 1 $ syntype. E. Peru, Whitely, 69-61. 
[PL 5, fig- 54-] 

Currently placed as a subspecies of Eueides isabella (Cramer) . 

hippolyte Bates, 1862 : 559 (as sp. of Heliconius). 1 <3, 1 $ syntypes. Godman-Salvin Coll. 
1913-2. 1 q\ Tapajos, Amazons. Bates. 1 $, Serpa, Amazons. Bates. [PL 24, fig. 241.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene madeira Riley. 

humboldti Neustetter, 1928a : 442 (as sp. of Heliconius). 1 o*. 1 $ syntypes. Rio Itaya by 
Iquitos. / Joicey Bequest. Brit. Mus. 1934-120. [PL 15, fig. 156.] 
Currently placed as a subspecies of Heliconius hecale (Fabricius). 

hydara Hewitson, 1867a : pi. 5, fig. 14 (as sp. of Heliconia). 1 q* syntype. N. Granada, 
Hewitson Coll. 79.69. [PL 31, fig. 316.] 

Currently placed as a subspecies of Heliconius erato (L.). 



TYPE-SPECIMENS OF HELICONIINAE i 93 

hydarina Stichel, 1912 : 1 (as form of Heliconius hermathena) . Holotype $. Faro. v. 1911 
Ducke. / N. Brazilien, Faro, e. c. H. Stichel. / Rothschild Bequest. B. M. 1939-1. [PI. 29, 
fig. 289.] 

Currently placed as a synonym of Heliconius hermathena veveatta Stichel. 

hygiana Hewitson, 1867a : pi. 5, fig. 15 (as sp. of Heliconia). 1 $ syntype. Quito, Hewitson 
Coll. 79.69. [PI. 36, fig. 365.] 

Currently placed as a subspecies of Heliconius clysonymus Latreille. 

ignotus Joicey & Kaye, igiyb : 89 (as sp. of Heliconius). 1 o* syntype. Charape, N. Peru. 
A. & E. Pratt. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 11, fig. 114.] 
Currently placed as a form of Heliconius numata talboti Joicey & Kaye. 

ilia Niepelt, 1908 : 506 (as form of Heliconius erato estrella). Holotype o*. Ob. Pastaza, 
Ecuad., c. 1000 m. Cll. Niepelt. / Adams Bequest. B. M. 1912-399. [PI. 30, fig. 307.] 

Currently placed as a transitional form between Heliconius erato notabilis Salvin & Godman 
and Heliconius erato lalivitta Butler. 

immoderata Stichel, 1906a : 9 (as form of Heliconius ismenius ismenius). 1 (J, 1 $ syntypes. 
Columb., Muzo. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 13, fig. i33l 

incarnata Riley, 1926 : 243 (as subsp. of Dione vanillae). Lectotype <J, 2 5 6*. x 3 $ paralecto- 
types. Lectotype <$, 1 $, 1 $ paralectotypes, Near Durango city, Mexico. Becker. / Godman- 
Salvin Coll. 191 5-3. 1 <J paralectotype, Florida. W. Schaus. 1905-244. 2 £, 2 $ paralecto- 
types, Florida, Morrison. / Godman-Salvin Coll. 191 5-3. 1 <J paralectotype, Florida, H. K. 
Morrison. / Godman-Salvin Coll. 1915-3. 1 o\ 1 $ paralectotypes, S. California: Pasadena, 
ix. 1920. G. B. Pearson. 1 <J paralectotype, San Diego. August. Cockerell. 1904-165. 1 o* 
paralectotype, Texas Hg. 71. / Zell. Coll. 1884. 1 q" paralectotype, Texas U.S.A. Belfrage. / 
Godman-Salvin Coll. 1915-3. 1 $ paralectotype, U.S. America: Havana 111. A. W. J. 
Pomeroy. 1919-269. / Pres. by Imp. Bur. Ent. 1 q" paralectotype, Mazatlan, Mexico. 
Forrer. / Godman-Salvin Coll. 1915-3. 1 q* paralectotype, Atoyac, Vera Cruz. May H. H. S. / 
Godman-Salvin Coll. 1915-3. 1 q* paralectotype, Cordova, Vera Cruz. Rumeli. / Godman- 
Salvin Coll. 1915-3. 1 <J paralectotype, Cordova, Mexico. Hoege. / Godman-Salvin Coll. 
1915-3. 1 (J paralectotype, Oaxaca, S. Mexico. Fenochio. / Godman-Salvin Coll. 1915-3. 2 <J, 
1 $ paralectotypes, Teapa, Tabasco. March. H. H. S. / Godman-Salvin Coll. 1915-3. 1^,1? 
paralectotype, Polochic Valley. F. D. G. & O. S. / Godman-Salvin Coll. 1915-3. 1 6* para- 
lectotype, Nicaragua. Sr. M. D. Chaves. 1900-65. (9). 20* paralectotypes, Matagalpa, 
Nicaragua. Richardson. / Godman-Salvin Coll. 1915-3. 1 <J paralectotype, Chontales, 
Nicaragua. Janson. / Godman-Salvin Coll. 1915-3. 2 q* paralectotypes, Costa Rica. Van 
Patten. Druce Coll. / Godman-Salvin Coll. 1915-3. 1 £ paralectotype, S. Fran'sco, 
4500 ft [1370 m] H. Rogers. / Godman-Salvin Coll. 1915-3. 1 9 paralectotype, 
Key West Fla. 30.1. '69. E. B. Zell. Coll. 1884. 1 <j> paralectotype, Texas Hg. 71. / Dallas 
Tex. Boll. / Zell. Coll. 1884. 1 $ paralectotype, Ent. Club. 44-12. 1 $ paralectotype, 
Alamos, Mexico. Buchan-Hepburn. / Godman-Salvin Coll. 1915-3. 1 $ paralectotype, Pinos 
Altos, Chihuahua, Mexico. Buchan-Hepburn. / Godman-Salvin Coll. 1915-3. 1 $ para- 
lectotype, S. Geronimo. Guatemala. Champion. / Godman-Salvin Coll. 1915-3. 1 $ para- 
lectotype, Honduras: Tegucigalpa i.xii.1920. B. M. 1921-503. [PI. 1, fig. 10.] 

Currently placed as a subspecies of Agraulis vanillae (L.). 

Lectotype designated by Michener (1942 : 7). 

indecisa Joicey & Kaye, igiyb : 91 (as subsp. of Heliconius aristiona). 1 <J syntype. Up. 
- Orinoco. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 15, fig. 153.] 
Currently placed as a form of Heliconius hecale ithaca Felder & Felder. 

insolita Riffarth, 19076 : 508 (as form of Heliconius melpomene timareta). 1 <$, 1 $ syntypes. 



194 P- R- ACKERY & R. L. SMILES 

Santa Jnez (Ecuad.). R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. 
Mus. 1934-120. [PL 28, fig. 285.] 

Currently placed as a form of Heliconius timareta Hewitson. 

insularis Stichel, 1909 : 179 (as subsp. of Eueides ricini). Holotype q\ Trinidad, Belmont, 
e. c. H. Stichel. / Rothschild Bequest. B. M. 1939-1. [PL 36, fig. 367.] 
Currently placed as a synonym of Heliconius ricini (L.) . 

intermedins Riffarth, 19076 : 509, pi. 5, fig. 11 (as subsp. of Heliconius hecuba). Holotype (J. 
Columbien. ex. coll. Fruhstorfer. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. 
Mus. 1934-120. [PL 7, fig. 75.] 

Currently placed as a transitional form between Heliconius hecuba hecuba Hewitson and 
Heliconius hecuba cassandra Felder & Felder. 

iris Riffarth, 19076 : 506 (as form of Heliconius melpomene aglaope). Holotype $. Ob. 
Pastaza, Ecuad., c. 1000 m. Cll. Niepelt. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PL 20, fig. 202.] 

Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene plesseni Riffarth. 

isolda Niepelt, 1908 : 505 (as form of Heliconius melpomene aglaope). 1 o* syntype. Jibaria. / 
Original. / Rothschild Bequest. B. M. 1939-1. [PL 20, fig. 203.] 

Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene plesseni Riffarth. 

isabellinus Bates, 1862 : 554 (as var. of Heliconius numata). 1 $, 1 $ syntypes. Godman- 
Salvin Coll. 191 3-2. 1 o*. S. Paulo, U. Amazons. H. W. Bates. 1 $, Tunantins, Amazons. 
H. W. Bates. [PL 11, fig. 113.] 

Currently placed as a subspecies of Heliconius numata (Cramer). 

ithaca Felder & Felder, 18626 : 418 (as sp. of Heliconius). 2 <J, 1 $ syntypes. Bogota, Lindig. / 
Felder Colin. / Rothschild Bequest. 1939-1. [PL 15, fig. 152.] 
Currently placed as a subspecies of Heliconius hecale (Fabricius). 

jucundus Bates, 1864a : 56 (as sp. of Heliconius). 10 (J, 3 $ syntypes. Lion Hill, Panama, 
McLeannan. / Godman-Salvin Coll. 1913-2. [PL 14, fig. 146.] 

Currently placed as a transitional form between Heliconius hecale melicerta Bates and 
Heliconius hecale zuleika Hewitson. 

juno Riffarth, 1900 : 209 (as var. of Heliconius cyrbia). Holotype o*- Original. / Ex. Coll. 
Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 31, fig. 317.] 

Probably best regarded as a synonym of Heliconius erato hydara form guarica Reakirt. 

kruegeri Neustetter, 1925 : 60 (as subsp. of Heliconius phyllis). Holotype <$. Rio Grande, 
Bolivia. / Joicey Bequest. Brit. Mus. 1934-120. [PL 34, fig. 344.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius). 

lampeto Bates, 1862 : 562 (as sp. of Eueides). Holotype $. S. Paulo, U. Amazons. H. W. 
Bates. / Godman-Salvin Coll. 1913-2. [PL 3, fig. 32.] 

lativitta Butler, 1877 : 150 (as sp. of Heliconius). 2 0*. x ? syntypes. 1 o*. Guayaquil, 
Stephens. 55-48. 1 q\ Boa vista, R. Madeira 1.6.74. / Amazons Trail. 77-64. / 77.65. Trail. 
1 $, Ega, Amazons, Bates. 56-69. [PL 31, fig. 311.] 

Currently placed as a subspecies of Heliconius erato (L.). 

latus Riffarth, 1900 : 197 (as var. oi Heliconius par aensis). 3 6*. 3 ? syntypes. Original. / Ex. 



TYPE-SPECIMENS OF HELICONIINAE 195 

Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 2 <J, 2 $, Itaituba, Stgr. 1 <$, Itait. 
1 9, Amaz. Sup. [PI. 16, fig. 166.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius). 

laurentina Neustetter, 19276 : 229 (as form- of Heliconius melpomene). Holotype <$. St. 
Laurent, Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 235.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene thelxiope (Hiibner). 

lavinia Neustetter, 1926& : 290 (as form of Heliconius melpomene) . 1 <$, 1 $ syntypes. St. 
Laurent. Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 22, fig. 226.] 

Currently placed as a transitional form between Heliconius melpomene meriana Turner and 
Heliconius melpomene melpomene (L.). 

leonora Kriiger, R., 1927 : 86 (as form of Heliconius phyllis). Holotype o". Rio Grande, 
Bolivia. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 34, fig. 343.] 

Currently placed as a transitional form between Heliconius erato venustus Salvin and 
Heliconius erato phyllis (Fabricius). 

lepidus Riffarth, 19076 : 503, pi. 5, fig. 2 (as subsp. of Heliconius aristiona). 2 <J, 3 $ syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 2 <J, 2 $, Coca, (Ecuad.), 
R. Haensch. S. 1 $, Sara-yacu (Ecuad.), R. Haensch. [PI. 10, fig. 103.] 
Currently placed as a form of Heliconius numata euphone Felder & Felder. 

leucadia Bates, 1862 : 556 (as sp. of Heliconius). Holotype o*- S. Paulo, U. Amazons. H. W. 
Bates. / Godman-Salvin Coll. 1913-2. [PI. 37, fig. 380.] 

leucomma Bates, 1866 : 88 (as sp. of Eueid.es). 3 <J, 1 ? syntypes. Lion Hill, Panama. 
McLeannan. / Godman-Salvin Coll. 1913-2. [PI. 6, fig. 64.] 

Currently placed as a synonym of Eueides lybia olympia (Fabricius). 

lindigii Felder & Felder, 1865 : 377, pi. 47, figs 1, 2 (as sp. of Heliconius). 1 <$ syntype. 
Bogota. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 8, fig. 89.] 
Currently placed as a subspecies of Heliconius burneyi (Hiibner). 

lineata Salvin & Godman, 1868 : 145 (as sp. of Eueides). 2 <J, 1 $ syntypes. Polochic Valley. 
F. D. G. & O. S. / Godman-Salvin Coll. 1913-2. [PI. 4, fig. 44.] 

longarena Hewitson, 1875 : 182 (as sp. of Heliconia). 1 q* syntype. N. Granada. Hewitson 
Coll. 79.69. [PL 35. fig- 357] 

Currently placed as a subspecies of Heliconius hecalesia Hewitson. 

lucia Riley, 1926 : 241 (as subsp. of Colaenis julia). 3 (J, 1 $ syntypes. 2 o\ 1 $, Santa Lucia. 
91-57. 1 (J, Sta. Lucia. W. Indies Comm. 1901-102. [PI. 2, fig. 22.] 
Currently placed as a subspecies of Dry as iulia (Fabricius). 

lucina Felder & Felder, 1862a : no (as sp. of Agraulis). 2 <J syntypes. Rio Negro. / Felder 
Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 2, fig. 12.] 
Currently placed as a subspecies of Agraulis vanillae (L.). 

lucindella Joicey & Kaye, 1917a : 423 (as ab. of Heliconius melpomene thelxiope). Holotype o*. 
Guyane Francse., St. -Jean du Maroni. Collection Le Moult. /Joicey Bequest. Brit. Mus. 
1934-120. [PI. 24, fig. 239.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene thelxiope (Hiibner). 

luminosus Riffarth, 1901 : 132 (as subsp. of Heliconius erato). 1 $, 1 $ syntypes. Columb. 
Stgr. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 9, fig. 93.] 
Currently placed as a form of Heliconius doris eratonius Staudinger. 
Described from Chiriqui. 



ig6 P. R. ACKERY & R. L. SMILES 

luteipicta Neustetter, 19266 : 281 (as form of Heliconius melpomene) . 1 $, 1 9 syntypes. 
St. Laurent, Guyana. / Joicey Bequest. Brit. Mus. 1934-120. 1 g, Co-Type. 1 9, Type. 
[PL 21, fig. 212.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

lutulenta Stichel, 1907 : 13 (as female form of Colaenis phaetnsa). Holotype 9- Paraguay, 
Nuev. Germania, e. c. H. Stichel. / Typus / Rothschild Bequest. B. M. 1939-1. [PI. 1, 

fig- 5-] 

Currently placed as a form of Dryadula phaetnsa (L.). 

maculosa Stichel, 1907 : 18 (as subsp. of Dione vanillae). 2 q\ 2 9 syntypes. Typus. 1 <£, 

1 $, Brazilia, Espirito Santo, e. c. H. Stichel. / Rothschild Bequest. B. M. 1 939-1. 1 5, 
Sud-Brazilien, Theresopolis. e. c. H. Stichel. / Rothschild Bequest. B. M. 1939-1. 1 <J, 
Paraguay, Nuev. Germania. e. c. H. Stichel. [PI. 2, fig. 11.] 

Currently placed as a subspecies of Agraulis vanillae (L.). 

madeira Riley, 1919 : 185 (as subsp. of Heliconius melpomene). 1 o" syntype. Lower Madeira, 
13. July 1913. / E. H. W. Wickham 1913-446. [PI. 24, fig. 244.] 

magdalena Bates, 1864a : 57 (as sp. of Heliconius). 1 5* syntype. Bogota, Colombia. Bates 
Coll. / Godman-Salvin Coll. 1913-2. [PI. 37, fig. 377.] 

Currently placed as a synonym of Heliconius sara sara (Fabricius) . 

majestica Joicey & Kaye, 1917a : 423 (as ab. of Heliconius melpomene thelxiope). Holotype o*- 
Guyane Francse., Nouveau Chantier. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 23, fig. 238.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) 
and Heliconius melpomene thelxiope (Hiibner). 

margarita Riffarth, 1900 : 205 (as ab. of Heliconius melpomene). 1 o" syntype. Bolivia. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 251.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 

maris Joicey & Kaye, 1917a : 418 (as ab. of Heliconius melpomene cybele). Holotype 9. 

2 o*. ! $ paratypes. Guyane Francse., St. Jean du Maroni. Collection Le Moult. / Joicey 
Bequest. Brit. Mus. 1934-120. [PI. 22, fig. 228.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

melanopors Joicey & Kaye, 1917a : 425 (as form of Heliconius numata). 2 $ syntypes. 
Joicey Bequest. Brit. Mus. 1934-120. 1 9, Guyane Francse., Nouveau Chantier. Collection Le 
Moult. 1 9. Guyane Francse., St. Jean du Maroni. Collection Le Moult. [PI. 12, fig. 124.] 
Currently placed as a form of Heliconius numata numata (Cramer). 

melete Felder & Felder, 1865 : 376 (as sp. of Heliconius). 2 (J, 1 9 syntypes. Nova Granada, 
Lindig. / Felder Colin. / Rothschild Bequest. B. M. 1 939-1. [PI. 8, fig. 83.] 
Currently placed as a subspecies of Heliconius xanthocles Bates. 

melicerta Bates, 1866 : 87 (as sp. of Heliconius) . 3 o* syntypes. Godman-Salvin Coll. 1913-2. 
2 q\ Lion Hill, Panama. McLeannan. 1 q*. Sta. Marta, N. Colombia. Bouchard. [PI. 14, 

fig- M3] 

Currently placed as a subspecies of Heliconius hecale (Fabricius). 

meliorina Neustetter, 19286 : 79 (as form of Heliconius estrella). 1^,1$ syntypes. S. 
Columbien, Mocoa. 1927. / Joicey Bequest. Brit. Mus. 1934-120. 1 o*> November. 1 9. 
Oktober. [PI. 31, fig. 313.] 



TYPE-SPECIMENS OF HELICONIINAE 197 

Currently placed as a. transitional form between Heliconius erato dignus Stichel and 
Heliconius erato lativitta Butler. 

tnellosa Stichel, 19066 : 208 (as subsp. of Colaenis euchroia). Lectotype $. Ecuador, 
e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. [PI. 2, fig. 16.] 
Currently placed as a subspecies of Podotricha euchroia (Doubleday). 
Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 615. 

melpina Joicey & Kaye, 1917a : 414 (as ab. of Heliconius melpomene melpomene) . Holotype $. 
Guyane Francse., St. Laurent du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 21, fig. 217.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

melpophylla Joicey & Kaye, 1917ft : 92 (as subsp. of Heliconius melpomene). Holotype $. 
Joicey Bequest. Brit. Mus. 1934-120. [PI. 19, fig. 192.] 

Currently placed as a transitional form between Heliconius melpomene amaryllis Felder & 
Felder and Heliconius melpomene aglaope Felder & Felder. 

messene Felder & Felder, 1862ft : 418 (as sp. of Heliconius). 1 o", 4 $ syntypes. Bogota. 
Lindig. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 9, fig. 101.] 
Currently placed as a subspecies of Heliconius numata (Cramer). 

metalilis Butler, 1873a : 167 (as sp. of Heliconius). 2 o*. l ? syntypes. Venezuela. Dyson. 
47-9. [PI. 17, fig. 169.] 

Currently placed as a subspecies of Heliconius ethilla Godart. 

mimulinus Butler, 1873a : 168 (as sp. of Heliconius). 2 <$, 1 $ syntypes. Bogota. Stevens. 
56-142. [PI. 8, fig. 84.] 

Probably best regarded as a synonym of Heliconius wallacei wallacei Reakirt. 

miraculosa Hering, 1926 : 199 (as sp. of Dione). 1 o* paratype. Arequipd, Siid Peru. leg. 
W. Hopp. / Presented by J. J. Joicey Esq., Brit. Mus. 1931-291. [PI. 1, fig. 6.] 
Currently placed as a subspecies of Dione juno (Cramer) . 

mirificus Stichel, 1906a : 12 (as subsp. of Heliconius silvana). Holotype $>. Ucayali. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. n, fig. 117] 
Probably best regarded as a synonym of Heliconius numata illustris Weymer. 

modesta Riffarth, 1900 : 206 (as ab. of Heliconius vulcanus). 2 (J, 1 $ syntypes. Paramba, 
Ecuador. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit, Mus. 1934-120. [PI. 18, 
fig. 181.] 

Currently placed as a transitional form between Heliconius melpomene cythera Hewitson 
and Heliconius melpomene vulcanus Butler. 

molina Grose-Smith, 1898 : 70 (as sp. of Heliconius). 1 <$ syntype. Valdevia, Colombia. 
Pratt, 97. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, fig. 295.] 
Currently placed as a transitional form between Heliconius erato chestertonii Hewitson and 
Heliconius erato hydara Hewitson. 

montanus Salvin, 1871 : 414 (assp. of Heliconius). 1 <J syntype. Orosi, Costa Rica. Kramer. / 
Godman-Salvin Coll. 1913-2. [PI. 35, fig. 361.] 

Currently placed as a subspecies of Heliconius clysonymus Latreille. 

mutabilis Butler, 1877 : 151 (as sp. of Heliconius) . 2 $ syntypes. 1 9. Serpa bush. 22.4.74. / 
Amazons Trail. 77-64. 1 $, Serpa, Amazons. Bates. / Godman-Salvin Coll. 1913-2. [PI. 23, 
fig. 230.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 



198 P. R. ACKERY & R. L. SMILES 

nanna Stichel, 18996 : 143 (as sp. of Heliconius). 2 q\ i ? syntypes. Brasilia, Espirito Santo, 
e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. [PI. 26, fig. 259.] 
Currently placed as a subspecies of Heliconius melpomene (L.). 

nattereri Felder & Felder, 1865 : 275, pi. 47, fig. 8 (as sp. of Heliconius). 1 <£ syntype. Bahia, 
Beske. / Felder Colin. / Rothschild Bequest. B. M. 1 939-1. [PI. 9, fig. 99.] 

The original type-series is clearly syntypic. The specimen included as holotype by 
Holzinger & Holzinger (1974) is merely an additional syntype. 

negroida Joicey & Kaye, 1917a : 418, pi. 107, fig. 2 (as ab. of Heliconius melpomene cybele). 
Holotype o*- Guyane Francse., St-Jean du Maroni. Collection Le Moult. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 22, fig. 222.] 

Currently placed as a synonym of Heliconius melpomene meriana form faustina Staudinger. 

negroidens Joicey & Kaye, 1917a : 424 (as ab. of Heliconius melpomene thelxiope?). 1 <J 
syntype. Guyane Francse., St-Jean du Maroni. Collection Le Moult. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 23, fig. 234.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene thelxiope (Hiibner). 

neustetteri Riffarth, 1908 : 1 14 (as subsp. of Heliconius alithea). 3 <$, 5 $ syntypes. Original. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 2 <J, 5 ?, Balzapamba (Ecuad.). 
R. Haensch S. 1 o". Ecuador, Rio Bamba. [PI. 28, fig. 283.] 
Currently placed as a form of Heliconius cydno alithea Hewitson. 

niepelti Riffarth, 19076 : 507, pi. 5, fig. 7 (as form of Heliconius melpomene aglaope). 1 <$ 
syntype. Pastaza sup., Ecuad. Oct-Dec. 06. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 20, fig. 201.] 

Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene plesseni Riffarth. 

nivea Kaye, 1916 : 195 (as ab. of Heliconius telesiphe). Lectotype <J- Perou, Collection Le. 
Moult. / Adams Bequest. B.M. 1912-399. [PL 35, fig. 359.] 

Currently placed as a synonym of Heliconius telesiphe cretacea Neustetter. 
Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 621. 

noctis Neustetter, 1926a : 36 (as form of Heliconius penelope). 1 £ syntype. Bolivia. 1925. 
W. Niepelt. Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 249.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene amandus Grose-Smith & Kirby. 

nocturna Riffarth, 1900 : 210 (as ab. of Heliconius hydara). Holotype <$. Venezuela, 
Fruhstorfer. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 29, 
fig. 296.] 

Currently placed as a transitional form between Heliconius erato chestertonii Hewitson and 
Heliconius erato hydara Hewitson. 

notabilis Salvin & Godman, 1868 : 145 (as sp. of Heliconius). 1 £ syntype. Canelos, Ecuador. 
Pearce. / Godman-Salvin Coll. 1913-2. [PI. 30, fig. 301.] 
Currently placed as a subspecies of Heliconius erato (L.). 
Described from female specimens only; however, this male is apparently a syntype. 

novatus Bates, 1867 : 539 (as sp. of Heliconius). 3 $ syntypes. Godman-Salvin Coll. 1913-2. 
1 $, Para. L. Amazons. H. W. Bates. 1 ?, Para. Bates. 49-49. 1 ?, Maranham, N. Brazil. 
T. Belt. [PI. 16, fig. 163.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius) . 



TYPE-SPECIMENS OF HELICONIINAE 199 

nubifer Butler, 1875 : 224 (as sp. of Heliconius). Holotype?. Fonteboa, Amazons. Bates. 
57-125. [PI. 10, fig. 112.] 

Currently placed as a subspecies of Heliconius numata (Cramer). 

numata Cramer, [1780] : 17, pi. 297, figs C, D (as sp. of Papilio). 1 <J syntype. Surinam. 
Cll. v. d. Capel. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 39, fig. 403.] 
Currently placed as a species of Heliconius Kluk. 

oberthueri Riffarth, 1902 : 162 (as subsp. of Heliconius erato). 2 $ syntypes. 1 ?, Berg en 
Daal, Sur., Juli. Michaelis. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. 1 $, Cayenne. / Levick Bequest. 1941-83. [PI. 32, fig. 326.] 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara (Hewitson). 

obscurata Riffarth, 1907b : 505 (as form of Heliconius melpomene funebris). Holotype $. 
Berg en Daal, Juli. Michaelis. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 22, fig. 220.] 

Currently placed as a form of Heliconius melpomene meriana Turner. 

obscurifascia Talbot, 1928 : 220 (as form of Heliconius melpomene burchelli). Holotype $. 
Melguira, 10 miles [16 km] S. of Diamantino. 2000 ft [610 m] 23. v. -3. vi. 27. Matto Grosso. 
C. L. Collenette. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 25, fig. 258.] 

Currently placed as a transitional form between Heliconius melpomene penelope Staudinger 
and Heliconius melpomene burchelli Poulton. 

obscurior Stichel, 1906a : 15 (as subsp. of Heliconius novatus). 1^,1 ? syntypes. Bolivia, 
Titicaca See. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 11. 
fig- H9-] 

Currently placed as a transitional form between Heliconius numata leopardus Weymer and 
Heliconius numata spadicarius Weeks. 

Occident ali s Neustetter, 1928^ : 258 (as subsp. of Heliconius ismenius). 1 $, 1 $ syntypes. 
Cantinela, Putamayo. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 13, fig. 132.] 
Currently placed as a form of Heliconius ismenius Latreille. 

ochracea Riffarth, 19076 : 511, pi. 5, fig. 12 (as form of Heliconius erato estrella). Holotype <$. 
Ob. Pastaza, Ecuad., c. 1000 m. Cll. Niepelt. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 30, fig. 306.] 

Currently placed as a transitional form between Heliconius erato notabilis Salvin & Godman 
and Heliconius erato lativitta Butler. 

octavia Bates, 1866 : 86 (as sp. of Heliconius). 2 (J syntypes. Duenas, Guatemala. F. D. 
G. & O. S. / Godman-Salvin Coll. 1913-2. [PI. 35, fig. 354.] 

Currently placed as a subspecies of Heliconius hecalesia Hewitson. 

pachinus Salvin, 1871 : 414 (as sp. of Eueides). 3 <J syntypes. Chiriqui, Panama. Arce. / 
Godman-Salvin Coll. 1913-2. [PI. 28, fig. 287.] 

pallens Stichel, 1903 : 19 (as form of Eueides vibilia vibilia). Holotype $. Espirito- 
Santo, Brazil, ex. coll. Fruhstorfer. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. 
Mus. 1934-120. [PI. 4, fig. 41.] 

pallida Riffarth, 19076 : 513 (as form of Eueides lampeto fuliginosus) . Holotype $. Ob. 
Pastaza, Ecuad., c. 1000 m. Cll. Niepelt. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 4, fig. 37.] 

Currently placed as an aberration of Eueides lampeto acacetes Hewitson. 

palmata Stichel, 1906a : 46 (as form of Heliconius erato estrella). Holotype $. Ucayali. / 



aoo P. R. ACKERY & R. L. SMILES 

Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 30, fig. 310.] 
Currently placed as a form of Heliconius erato lativitta Butler. 

paraensis Riffarth, 1900 : 197 (as sp. of Heliconius). 2 6* syntypes. Original. / Ex. Coll. 
Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 $, Para. 6.6.94. J 6*. Para 7.10.94. 
[PI. 16, fig. 167.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius). 

paraplesius Bates, 1867 : 540 (as sp. of Heliconius). 1 <$ syntype. Maranham, N. Brazil. 
T. Belt. / Godman-Salvin Coll. 1913-2. [PI. 8, fig. 81.] 

Currently placed as a subspecies of Heliconius xanthocles Bates. 

pardalinus Bates, 1862 : 555 (as var. of Heliconius eucoma). 2 <$ syntypes. S. Paulo, U. 
Amazons. H. W. Bates. / Godman-Salvin Coll. 1913-2. [PI. 13, fig. 137.] 
Currently placed as a species of Heliconius Kluk. 

parva Neustetter, 19286 : 77 (as form of Heliconius melpomene aglaope). Holotype 6*. S. 
Colombien, Mocoa. Oktober 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 20, fig. 207.] 
Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene bellula Stichel. 

parvimaculata Riffarth, 1900 : 207 (as ab. of Heliconius clytia). 8 (J, 3 $ syntypes. Original. / 
Joicey Bequest. Brit. Mus. 1934-120. 1 <£, Amazonas, Heyne. / Ex. Coll. Riffarth. 1 <£, Pari- 
catuba, 16.6.93. 6 $, 3 $, Obidos, Amazonen strom. VIII-IX.99. ex. coll. H. Fruhstorfer. / 
Ex. Coll. Riffarth. [PI. 8, fig. 87.] 

Currently placed as a form of Heliconius wallacei wallacei Reakirt. 

pasithoe Cramer, [1775] : 25, pi. 17, figs A, B (as sp. of Papilio). 3 $ syntypes. Felder 
Colin. / Rothschild Bequest. B. M. 1939-1. 1 <J, No. 2. Pasithoe. Cr., I, 17, A, B. / Surinam 
Coll. Linep. r o", Eyndhoven. 1 $, Coll. Linep. [PL 39, figs 400, 401, 402.] 
Currently placed as a synonym of Heliconius hecale hecale (Fabricius). 

paula Neustetter,- 19286 : 77 (as form of Heliconius melpomene aglaope). Holotype <J. S. 
Columbien, Mocoa. Oktober 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 20, fig. 208.] 

Currently placed as a transitional form between Heliconius melpomene aglaope Felder & 
Felder and Heliconius melpomene bellula Stichel. 

peeblesi Joicey & Talbot, 1925 : 647 (as sp. of Heliconius). 1 6", 1$ syntypes. MeridaDist., 
Venezuela. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 11, fig. 120.] 
Currently placed as a subspecies of Heliconius numata (Cramer). 

pellucida Srnka, 1885 : 130, pi. 1, fig. 3 (as sp. of Eueides). Holotype $. Pellucida Type 
Srnka. / Coll. Srnka assmann Cundinamarca. / Ex. Oberthiir Coll. Brit. Mus. 1927-3. [PI. 5, 

fig- 49-] 

Currently placed as a synonym of Eueides isabella huebneri Menetries. 

penelopides Neustetter, 19276 : 229 (as form of Heliconius melpomene). Holotype o". St. 
Laurent, Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 23, fig. 233.] 

Currently placed as a transitional form between Heliconius melpomene thelxiope (Hiibner) 
and Heliconius melpomene melpomene (L.). 

The original description is clearly based on one specimen only and the above specimen 
appears to be the true holotype. Holzinger & Holzinger (1974) also doubtfully include a 
specimen in the NM, Vienna as the 'Holotype' of penelopides. 

perchlora Joicey & Kaye, 191 76 : 94 (as subsp. of Heliconius elevatus). Holotype $. Boliv. / 
Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 17, fig. 178.] 

petiverana Doubleday, 1847 : 103 (as sp. of Heliconia). 1 $ syntype. Ex. Musaeo Boisduval. / 
Mexico. / Levick Bequest. 1941-83. [PI. 29, fig. 292.] 
Currently placed as a subspecies of Heliconius erato (L.). 



TYPE-SPECIMENS OF HELICONIINAE 201 

philipi Brown, 1975 : 214 (as subsp. of Heliconius aoede). Holotype $. Salampioni, Bolivia. 
Rosenb. / Ex. Coll. Herbert Druce, 1913. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 38, 
fig- 390.] 

phyllidis Grose-Smith & Kirby, 1892 : 2, pi. 1, fig. 4 (as sp. of Heliconius). 1 <J syntype. 
Boliv. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 34, fig. 352.] 
Currently placed as a form of Heliconius erato phyllis (Fabricius). 

phyllis Fabricius, 1775 : 463 (as sp. of Papilio). 1 <J syntype. Papilio Phyllis Fab. Entom. 
p. 463, n. 86. (In coll. J. Banks.) [PI. 34, fig. 349.] 
Currently placed as a subspecies of Heliconius erato (L.). 

plesseni Riffarth, 1907a : 333 (as form of Heliconius batesi). 1 <J syntype. Baraneus, Pastazza 
super. Ecuador. / Ex. Coll. Riffarth. / Original. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 19, fig. 193.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 

praelautus Stichel, 1906a : 10 (as subsp. of Heliconius numatus). Holotype $. Ucayali. / 
Original. / Ex. Coll. Biffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 10, fig. 109.] 
Currently placed as a female form of Heliconius numata arcuella Druce. 

pratti Joicey & Kaye, igiyb : 90, pi. 5, fig. 4 (as subsp. of Heliconius aristiona). 4 $ syntypes. 
Charape, N. Peru. 4000 ft [1220 m] A. E. & F. Pratt. / Joicey Bequest. Brit. Mus. 1934-120. 
3 ?, Sep.-Oct. 1912. 1 $, 1912. [PI. 10, fig. 107.] 

Currently placed as a form of Heliconius numata aristiona Hewitson. 

primularis Butler, 1869 : 18, pi. 9, fig. 2 (as sp. of Heliconius). 1 o". * ? syntypes. 1 £, 
Guayaquil. Stevens. 55-48. 1 $, Rio Napo. Stevens. 51-70. [Pi. 38, fig. 387.] 
Currently placed as a subspecies of Heliconius eleuchia Hewitson. 

primus Joicey & Kaye, 1917a : 414 (as ab. of Heliconius melpomene melpomene). Holotype $. 
Guyane Francse., St. Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 21, fig. 214.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene meriana Turner. 

problemata Neustetter, 19286 : 78 (as form of Heliconius erato estrella). 1 3* syntype. S. 
Columbien, Mocoa. November 1927. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 31, fig. 314.] 
Currently placed as a transitional form between Heliconius erato dignus Stichel and Heli- 
conius erato lativitta Butler. 

procula Doubleday, 1848 : 146, pi 20, fig. 1 (as sp. of Eueides). 1 o* syntype. Venezuela. 
Dyson. 46-75. [PI. 3, fig. 30.] 

protea Joicey & Kaye, 1917a : 427 (as ab. of Heliconius erato). Holotype $. Guyane Francse., 
St. Jean du Maroni, Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 33, 
fig- 332-] 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara Hewitson. 

punctarius Joicey & Kaye, 1917a : 422 (as ab. of Heliconius melpomene thelxiope). Holotype 
$. Guyane Francse., St. Jean du Maroni. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 24, 
fig. 240.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene thelxiope (Hiibner). 

punctata Hall, 1936 : 276 (as subsp. of Heliconius charithonia) . 7 q", 7 $ syntypes. 6 6*. 7 ?. 



202 P. R. ACKERY & R. L. SMILES 

St. Kitts, B. W. I. December 1935. A. Hall. / Brit. Mus. 1936-736. 1 <$, Antigua. J. W. 
Gregory. 99-164. [PI. 36, fig. 366.] 

Currently placed as a subspecies of Heliconius charitonia (L.). 

pur a Niepelt, 19076 : 42 (as form of Heliconius batesi plesseni). 2 <J, 3 ? syn types. Original. 
1 o*, Ob. Pastaza, Ecuad. c. 1000 m. Cll. Niepelt. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. 
Mus. 1934-120. 1 <$, 1 $, R. Pastaza. / Rothschild Bequest. B. M. 1939-1. 1 $, Ob. Pastaza, 
Ecuad., c. 1000 m. Cll. Niepelt. / Rothschild Bequest. B. M. 1939-1. 1 $, Ex. Coll. Riffarth. / 
Joicey Bequest. Brit. Mus. 1934-120. [PI. 19, fig. 196.] 

Currently placed as a form of Heliconius melpomene plesseni Riffarth. 

pygmalion Fruhstorfer, 1912 : 14 (as subsp. of Metamandana dido). Holotype $. Obidos. 
Fruhstorfer Coll. B. M. 1937-285. [PI. 1, fig. 1.] 
Currently placed as a species of Philaethria Billberg. 

quirina Cramer, [1775] : 101, pi. 65, figs A, B (as sp. of Papilio). 1 <J, 1 $ syntypes. Coll. 
Linep. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 39, figs 397, 398.] 
Currently placed as a synonym of Heliconius doris (L.). 

quitalena Hewitson, [1853] : pi. 1, fig. 3 (as sp. of Heliconia). 3 q* syntypes. 2 q\ Quito. 
Ffewitson Coll. 79.69. 1 o\ Ecuador. Hewitson Coll. 79.69. [Pi. 15, fig. 155.] 
Currently placed as a subspecies of Heliconius hecale (Fabricius). 

radiata Oberthiir, 1916 : 33, pi. 404, fig. 3479 (as form of Heliconius microclea notabilis). 
Holotype $. Guayaquil. / Levick Bequest. B. M. 1941-83. [Pi. 30, fig. 303.] 

Currently placed as a transitional form between Heliconius erato notabilis Salvin & Godman 
and Heliconius erato lativitta Butler. 

radiosus Butler, 1873a : 166 (as sp. of Heliconius). 1 $ syntype. Villa Nova, Amazons. 
Bates. 55-44- [ pl - J 6, fig- 165.] 

Currently placed as a subspecies of Heliconius pardalinus Bates. 

rhea Cramer, [1775] : 85, pl. 54, figs C, D (as sp. of Papilio). 1 q* syntype. Surin. Coll. 
Linep. / No. 8. rhea. Cr., 54, C, D. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. 
[Pl. 39, fig. 399.] 

Currently placed as a synonym of Heliconius sara thamar (Hiibner). 

rifjarthi Stichel, 1903 : 31, pl. 1, fig. 25 (as ab. of Eueides eanes). 2 6* syntypes. 1 <$, 
Stid-Peru, Chanchamayo. e. c. H. Stichel. / Typus. 1 o", Siid-Peru, Pozuzo. e. c. H. Stichel. / 
Typus. / Rothschild Bequest. B. M. 1939-1. [Pl. 4, fig. 46.] 

Currently placed as a transitional form between Eueides eanes heliconioides Felder & 
Felder and Eueides eanes pluto Stichel. 

riffarthi Stichel, 1906a : 26, pl. 2, fig. 8a (as subsp. of Heliconius melpomene). 2 $ syntypes. 
Ucayali. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 19, 
fig. 189.] 

Currently placed as a transitional form between Heliconius melpomene xenoclea Hewitson 
and Heliconius melpomene aglaope Felder & Felder. 

rondonia Brown, 1973 : 2 (as subsp. of Heliconius astraea). 5 o* paratypes. River System 
Cuyaba-Corumba, Matto Grosso, Brazil. / Joicey Bequest. Brit. Mus. 1934-120. [Pl. 9, 
fig. 92.] 

roraima Turner, 1966a : 109 (as subsp. of Heliconius elevatus). Holotype $, 3 <$, 1 $ paratypes. 
Holotype o*. 1 c?> x ? paratypes, Roraima, B. Guiana. H. Whitely. / Godman-Salvin Coll. 
1913-2. 1 o* paratype, PRoraima. Crowley Bequest. 1901-78. 1 q* paratype, Roraima, Br. 
Guiana. Crowley Bequest. 1901-78. [Pl. 17, fig. 175.] 

rosacea Riffarth, 19076 : 512, pl. 5, fig. 14 (as form of Heliconius erato simplex). Holotype o*. 



TYPE-SPECIMENS OF HELICONIINAE 



203 



Pastaza sup., Ecuador. Oct-Dez. 06. / Original. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 30, fig. 302.] 

Currently placed as a form of Heliconius erato notabilis Salvin & Godman. 

roseoflava Neustetter, 19266 : 292 (as form of Heliconius erato). 2 $ syntypes. St. Laurent, 
Guyana. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 33, fig. 333.] 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara Hewitson. 

rosina Boisduval, 1870 : 29 (as sp. of Heliconia). 1 5* syntype. Mexiq. / Ex. Musaeo Bois- 
duval. / Levick Bequest. 1941-83. [PI. 18, fig. 179.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 

rothschildi Niepelt, 1909 : 107 (as form of Heliconius erato estrella). 1 q*, 1 $ syntypes. 
Ecuador, Canelos. 800 m. / Rothschild Bequest. B. M. 1939-1. [PI. 30, fig. 305.] 

Currently placed as a transitional form between Heliconius erato notabilis Salvin & Godman 
and Heliconius erato lativitta Butler. 

rubellius Grose-Smith & Kirby, 1892 : 1, pi. 1, figs 1, 2 (as sp. of Heliconius). 1 <$ syntype. 
Columb. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 28, fig. 279.] 

Currently placed as a hybrid between Heliconius melpomene melpomene (L.) and Heliconius 
cydno cydno Doubleday. 

rubescens Stichel, 1906a : 26 (as form of Heliconius melpomene riffarthi). Holotype o*- 
Chuchurras, Peru. 320 m. Hoffmann. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. 
Mus. 1934-120. [PI. 19, fig. 190.] 

Currently placed as a transitional form between Heliconius melpomene xenoclea Hewitson 
and Heliconius melpomene aglaope Felder & Felder. 

rubicunda Niepelt, 1907& : 42 (as form of Heliconius batesi plesseni). 1 <J, 1 ? syntypes. 
Original. 1 q", Rothschild Bequest. B. M. 1939-1. 1 $, Ex. Coll. Riffarth. / Joicey Bequest. 
Brit. Mus. 1934-120. [PI. 19, fig. 194.] 

Currently placed as a form of Heliconius melpomene plesseni Riffarth. 

rubra Stichel, 1906a : 26 (as form of Heliconius melpomene aglaope). 2 o* syntypes. Original. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q", Cuzco. i £, Marcapata, 
Cuzco, Peru. [PI. 20, fig. 205.] 

Probably best regarded as a synonym of Heliconius melpomene aglaope Felder & Felder. 

rubrizona Joicey & Kaye, 191 jb : 93, pi. 4, fig. 7 (as subsp. of Heliconius erato). 1 6* syntype. 
Santarem. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 31, fig. 319.] 
Currently placed as a form of Heliconius erato hydara Hewitson. 

rufolimbata Butler, 1873a : 169 (as sp. of Heliconius). 1 o" syntype. Rio Tapayos, Brazil. 
Bates. 53-27. [PL 24, fig. 242.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L.) and 
Heliconius melpomene madeira Riley. 

schmassmanni Joicey & Talbot, 1925 : 647 (as subsp. of Heliconius elevatus). Holotype o*- 
River System Cuyaba-Corumba, Matto Grosso, Brazil. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 17, fig. 177.] 

schulzi Riffarth, 1899 : 405 (as sp. of Heliconius) . 2 $ syntypes. Original. / Ex. Coll. Riffarth. / 
Joicey Bequest. Brit. Mus. 1934-120. 1 $, Para, 21.12.93. A. Schulz. 1 $, Para, 2.93. A. 
Schulz. [PI. 16, fig. 164.] 

Currently placed as a synonym of Heliconius hecale novatus Bates. 

seitzi Stichel, 1903 : 6, pi. 1, fig. 6 (as subsp. of Eueides isabella). 1 o* syntype. Santa Jnez. 



2o 4 P. R. ACKERY & R. L. SMILES 

(Ecuad.) R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 5, fig- 52.] 

Currently placed as a form of Eueides Isabella dissoluta Stichel. 

semiphorus Staudinger, 1896 : 284 (as var. of Heliconius metaphorus). 1 <J syntype. Rio 
Dagua. Stgr. / Origin. / Ex. Coll. Riffarth. [PL 14, fig. 145.] 
Currently placed as a form of Heliconius hecale melicerta Bates. 

semirubra Joicey & Kaye, 191 jb : 90 (as ab. of Heliconius clysonimus apicalis). 1 <J syntype. 
Siato, Rio Siato, Slopes of Choco, Columbia. 5200 ft [1580 m] Sept. '09. / Ex. Coll. Herbert 
Druce. 1913. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 36, fig. 363.] 

Currently placed as a transitional form between Heliconius clysonymus clysonymus Latreille 
and Heliconius clysonymus hygiana Hewitson. 

seraphini Talbot, 1932 : 191 (as sp. of Heliconius). Holotype 5". Guyane, St. Laurent. 
Joicey Bequest. Brit. Mus. 1934-120. [PI. 12, fig. 123.] 

Currently placed as a hybrid between Heliconius numata numata (Cramer) and Heliconius 
melpomene thelxiope (Hiibner). 

serpensis Kaye, 1919 : 218 (as subsp. of Heliconius burneyi). 1 <J, 1 $ paratypes. Serpa, 
Lower Amazon. Jan-March 1914. A. Hall. / Brit. Mus. 1933-61. [PI. 8, fig. 90.] 

Currently placed as a transitional form between Heliconius burneyi burneyi (Hiibner) and 
Heliconius burneyi catherinae Staudinger. 

silvana Stoll, [1781] : 143, pi. 366, figs C, D (as sp. oiPapilio). 1 $ syntype. No. 32. Silvana. 
Cr., IV, 364, C, D. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 39, fig. 404.] 
Currently placed as a subspecies of Heliconius numata (Cramer) . 

silvaniformis Joicey & Kaye, 191 76 : 89, pi. 5, fig. 7 (as subsp. of Heliconius numata). 3 0* 
syntypes. Para, Lower Amazon. Jan-March 1914. A. Hall. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 12, fig. 127.] 

Currently placed as a form of Heliconius numata superioris Butler. 

simplex Riffarth, 1906 : 56 (as form of Heliconius erato estrella). Holotype $. Cajon, Siid- 
Peru. Indeinw., Callao. / Original. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 33, fig. 336.] 
Currently placed as a transitional form between Heliconius erato lativitta Butler and 
Heliconius erato amphitrite Riffarth. 

sincerus Riffarth, 19076 : 501 (as subsp. of Heliconius numatus). i(J, 1$ syntypes. Original./ 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 5*, Amazon inf. 1 $, Venezuela. 
[PI. 12, fig. 125.] 

Currently placed as a form of Heliconius numata superioris Butler. 

sisyphus Salvin, 1871 : 413 (as sp. of Heliconius). 2 <$ syntypes. Cosnipata Vail. E. Peru. 
H. Whitely. / Godman-Salvin Coll. 1913-2. [PL 15, fig. 158.] 
Currently placed as a subspecies of Heliconius hecale (Fabricius) . 

sotericus Salvin, 1 871 : 414 (as sp. of Heliconius). Lectotype <J, 5 <J paralectotypes. Guaymay 
Ecuador. C. Buckley. / Godman-Salvin Coll. 1913-2. [PL 35, fig. 358.] 
Currently placed as a subspecies of Heliconius telesiphe Doubleday. 
Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 621. 

sperata Riffarth, 19076 : 513 (as form of Heliconius erato phyllis). 2 5*, 1 $ syntypes. Original. 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934—120. 1 o*> Cajon, Siid-Peru. Indienw., 
Callao. 1 o", 1 ?, Prov. Sara, Dep. St. Cruz, Siid Bolivia. [PL 34, fig. 350.] 

Currently placed as a transitional form between Heliconius erato phyllis (Fabricius) and 
Heliconius erato venustus Salvin. 



TYPE-SPECIMENS OF HELICONIINAE 205 

spoliata Stichel, 1903 : 9 (as ab. of Heliconius Isabella huebneri). Holotype $. S. America, 
Caucathal. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 5, 

fig- 51J 

Currently placed as a form of Eueides Isabella eva Fabricius. 

sprucei Bates, 1864a : 57 (as sp. of Heliconius). 4 o*. l ? syntype. Chimborazo, Ecuador. 
Spruce. Bates Coll. / Godman-Salvin Coll. 1913-2. [PL 37, fig. 378.] 
Currently placed as a subspecies of Heliconius sara (Fabricius). 

sticheli Riffarth, 19076 : 509, pi. 5, fig. 10 (as subsp. of Heliconius vulcanus). 4 o\ 1 $ syntypes. 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 3 J 1 $, Paramba, 
Ecuador. 1 q\ Paramba, 3,500 ft II. 97. Dry Season. (Rosenberg). [PL 18, fig. 182.] 

Currently placed as a transitional form between Heliconius melpomene cythera Hewitson and 
Heliconius melpomene vulcanus Butler. 

straminea Riley, 1926 : 242 (as subsp. of Colaenis euchroia). Holotype q*- Balsapamba, 
Ecuador. / Crowley Bequest. 1901-78. [PL 2, fig. 17.] 

Currently placed as a subspecies of Podotricha euchroia (Doubleday). 

stupenda Stichel, 1907 : 13 (as form of Colaenis phaetusa). Holotype $. Panama, Bugaba. 
e. c. H. Stichel. /' Typus. / Rothschild Bequest. B. M. 1939-1. [PL 1, fig. 3.] 
Currently placed as a form of Dvyadula phaetusa (L.). 

stygianus Joicey & Kaye, 1917a : 423 (as ab. of Heliconius melpomene thelxiope). Holotype 
o*. Guyane Francse., St-Jean du Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 
1934-120. [PL 23, fig. 231.] 

Currently placed as a transitional form between Heliconius melpomene thelxiope (Hiibner) 
and Heliconius melpomene melpomene (L.). 

styx Niepelt, 192 1 : 19 (as form of Heliconius andevida). 2 £ syntypes. Joicey Bequest. Brit. 
Mus. 1934-120. 1 q\ Dep. Malacatan, Guatemala. [PL 13, fig. 138.] 

Currently placed as an aberrational form of Heliconius hecale fovnarina Hewitson. 

submarginatus Fassl, 191 2 : 56 (as subsp. of Heliconius weymeri). 3 o*. 3 ? syntypes. Rio 
Aguacatal, Colomb., W. Codr. 2000 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 27, fig. 275.] 

Currently placed as a form of Heliconius cydno weymeri Staudinger. 

subnubilis Stichel, 1906a : 14, pi. 1, fig. 4a (as subsp. of Heliconius novatus). 2 $ syntypes. 
Juanjuy, Peru. / Original. / Joicey Bequest. Brit. Mus. 1934-120. [PL 11, fig. 116.] 
Currently placed as a form of Heliconius numata illustris Weymer. 

sulphureomaculata Fassl, 1914 : 43 (as ab. of Heliconius weymeri). 1 $ syntype. Rio 
Aguacatal, Colomb., W. Cord. 2000 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. 
[PL 27, fig. 274.] 

Currently placed as a form of Heliconius cydno weymeri Staudinger. 

super ba Lathy, 1906 : 453, pi. 34, fig. 2 (as var. of Heliconius xenoclea). Holotype o*- Rio 
Colorado, Peru. 2500 ft [760 m] ix.03. Watkins & Tomlinson. / Adams Bequest. B. M. 1912-399. 
[PL 18, fig. 188.] 

Currently placed as a transitional form between Heliconius melpomene xenoclea Hewitson 
and Heliconius melpomene aglaope Felder & Felder. 

superioris Butler, 1875 : 224 (as sp. of Heliconius). 1 <$, 1 $ syntypes. 1 q". Villa Nova, 
Amazons. Bates. 55-75. 1 $, Ega, Braz. [PL 12, fig. 126.] 

Currently placed as a subspecies of Heliconius numata (Cramer) . 

surdus Stichel, 1903 : 27, pi. 1, fig. 20 (as subsp. of Heliconius thales). 1 o*. T 9 syntypes. 



206 P. R. ACKERY & R. L. SMILES 

Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q\ Demerara. 1 $, 
Cameta. 12.92. [PI. 5, fig. 58.] 

Currently placed as a form of Eueides tales tales (Cramer). 

tabaconas Brown, 1975 : 221 (as subsp. of Heliconius clysonymus). Holotype $, 1 £, 2 $ 
paratypes. Joicey Bequest. Brit. Mus. 1934-120. Holotype o*> Charape, N. Peru, 4000 ft 
[1220 m], Sep.-Oct. 1912. A. & E. Pratt. 1 <J, 1 $■ River Tabaconas, N. Peru, 6000 ft [1800 m] 
A. E. & F. Pratt. 1912. 1 $, Charape, N. Peru. 4000 ft [1220 m] A. E. & F. Pratt. [PI. 38, 
fig- 39i •] 

talboti Joicey & Kaye, 19176 : 88, pi. 5, fig. 3 (as subsp. of Heliconius numata). 5 $, 1 <j> 
syntypes. Joicey Bequest. Brit. Mus. 1934-120. 1 <$, River Chinchipe, N. Peru. 6000 ft 
[1800 m] Sep. 1912. A. & E. Pratt. 1 o*> Charape, N. Peru. 4000 ft [1220 m] Sept. -Oct. 
1912. A. & E. Pratt. 1 $, Rentema Falls, Upper Maranon, N. Peru. 1000 ft [300 m] A. & E. 
Pratt. 2 o\ 1 $. Charape, N. Peru. 4000. A. E. & F. Pratt. 1912. [PI. n, fig. 115.] 

tarapotensis Riffarth, 1901 : 59 (as subsp. of Heliconius aristiona). 1 q\ 2 $ syntypes 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 q*. Tarapota, Peru. 
Garlepp. 2 9, Yuan-yui, Peru. Garlepp. [PI. 10, fig. 105.] 

Currently placed as a form of Heliconius numata euphone Felder & Felder. 

Described from Tarapota. 

tecta Riffarth, 1900 : 207 (as ab. of Heliconius erato). 1 <J syntype. Columbien. ex. coll. 
Suffert. / Original. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 9, fig. 94.] 
Currently placed as a form of Heliconius doris dovis (L.). 

telchinia Doubleday, 1847 : pi. 14, fig. 4 (as sp. of Heliconia) . 1 $ syntype. Honduras. Purch. 
from Dyson. 45-123. [PI. 12, fig. 129.] 

Currently placed as a subspecies of Heliconius ismenius Latreille. 

telesiphe Doubleday, 1847 : 103, pi. 15, fig. 2 (as sp. of Heliconia). Lectotype <$, 1 <$ para- 
lectotype. Bolivia. Bridges. 46-76. [PI. 35, fig. 360.] 
Currently placed as a species of Heliconius Kluk. 
Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 620. 

telesiphe Hewitson, 1 867ft : 5^4 (as sp. of Colaenis). Lectotype <J, 2 $, 1 $ paralectotypes. 
Ecuador. Hewitson Coll. 79.69. [PI. 2, fig. 19.] 

Currently placed as a species of Podotricha Michener. 

Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 617. 

telloides Joicey & Talbot, 1925 : 648 (as ab. of Heliconius erato). Holotype $. Guyane 
Francaise, Godebert-Maroni. Collection Le Moult. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 33, fig. 331.] 

Currently placed as a transitional form between Heliconius erato erato (L.). and Heliconius 
erato hydara (Hewitson). 

tellus Oberthur, 1902 : pi. 3, fig. 26 (as var. of Heliconius vesta). Holotype q\ Cayenne. / 
Levick Bequest. 1941-83. [PI. 32, fig. 330.] 

Currently placed as a transitional form between Heliconius erato erato (L.) and Heliconius 
erato hydara Hewitson. 

temerinda Hewitson, 1873 : pi. 7, fig. 23 (as sp. of Heliconia). 1 5* syntype. Villagomes, 
N. Granada. / Hewitson Coll. 79.69. [PI. 26, fig. 268.] 

Currently placed as a form of Heliconius cydno hermogenes Hewitson. 

thetis Boisduval, 1870 : 29 (as sp. of Heliconia). Holotype $. Nicaragua. / Ex Musaeo 
Boisduval. / Levick Bequest. 1941-83. [PI. 6, fig. 70.] 

Currently placed as a subspecies of Heliconius metharme Erichson. 



TYPE-SPECIMENS OF HELICONIINAE 207 

theudela Hewitson, 1874 : 224 (as sp. of Heliconia). 1 o* syntype. Panama. Hewitson Coll. 
79.69. [PI. 37, fig. 376.] 

Currently placed as a subspecies of Heliconius sara (Fabricius). 

thyana Felder & Felder, i860 : 105 (as sp. of Eueides). 1 $ syntype. Brazil Mus. Type. / 
Rothschild Bequest. B. M. 1939-1. [PI. 4, fig. 43.] 

Currently placed as a female form of Eueides pavana Menetries. 

timareta Hewitson, 18676 : 563 (as sp. of Heliconia). 1 <J syntype. Ecuador. Hewitson 
Coll. 79.69. [PI. 28, fig. 286.] 

Currently placed as a species of Heliconius Kluk. 

tithraustes Salvin, 1871 : 415 (as sp. of Colaenis). Lectotype 6*. 4 6* paralectotypes 
Godman-Salvin Coll. 1915-3. Lectotype <J, 1 6* paralectotype, Rio Topo, Ecuador, C. Buckley 
Druce Coll. 1 $ paralectotype, Canelos, Ecuador. 1 6* paralectotype, Canelos, Ecuador, 
Pearce. 1 <J paralectotype, Santa Inez, Buckley. [PI. 2, fig. 18.] 

Currently placed as a subspecies of Podotricha telesiphe (Hewitson). 

Lectotype designated by Vane-Wright, Ackery & Smiles, 1975 : 618. 

titio Stichel, 1907 : 12 (as subsp. of Colaenis iulia). 2 o*, 1 $ syntypes. Typus. / Rothschild 
Bequest. B. M. 1939-1. 1^,1$, Sta. Cruz, Slid Bolivia. / e. c. H. Stichel. 1 <3\ Bolivia, 
Yungas d. 1. Paz. e. c. H. Stichel. [PI. 3, fig. 25.] 

Currently placed as a subspecies of Dryas iulia (Fabricius). 

tolima Fassl, 1912 : 55 (as sp. of Heliconius). 5 <$, 4 $ syntypes. 3 <J, 1 9, Canon del Tolima. 
Columb. Cent. Cord. 1700 m. Coll. Fassl. / Joicey Bequest. Brit. Mus. 1934-120. 1^,1$, 
Canon del Tolima. Columbien. 1700 m. Fassl. / Ex. Coll. Herbert Druce. 1913. / Joicey 
Bequest. Brit. Mus. 1934-120. 1 <$, Cavon del Tolima 7700 m. Dec. 09. / Levick Bequest. 
1941-83. 1 $, XL 09. Canon del Tolima. 1700 m. / Levick Bequest. 1941-83. 1 $, Canon 
del Mte Tolima, Colombia 1700 m. 12.09 / Adams Bequest. B. M. 1912-399. [PI. 7, fig. 73.] 
Currently placed as a subspecies of Heliconius hecuba Hewitson. 

translata Joicey & Kaye, 191 76 : 91 (as ab. of Heliconius numata superioris). 1 $ syntype. 
Manaos, Lower Amazon, Jan. -March 1914. A. Hall. / Joicey Bequest. Brit. Mus. 1934-120. 
[PI. 11, fig. 121.] 

Currently placed as a form of Heliconius numata mavors Weymer. 

tristis Riffarth, 1900 : 210 (as ab. of Heliconius petiver ana). Holotype o". Chiriqui. / Original. 
/ Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 31, fig. 315.] 

Currently placed as a transitional form between Heliconius erato hydara Hewitson and 
Heliconius erato petiverana Doubleday. 

turneri Brown & Benson, 1975 : 2 (as subsp. of Heliconius demeter). Holotype <$. Ypiranga, 
H. Boy. xi. 29. [PI. 36, fig. 373.] 

tyche Bates, 1862 : 559 (as sp. of Heliconius). 1 <? syntype. Serpa, Amazons. Bates. / 
Godman-Salvin coll. 1913-2. [PI. 24, fig. 243.] 

Currently placed as a transitional form between Heliconius melpomene melpomene (L) and 
Heliconius melpomene madeira Riley. 

umbratilis Rober, 1927 : 402 (as subsp. of Eueides edias). 2 o\ 1 ? syntypes. 1 6*. * ?. 
Sierra Nevada de Sta. Marta, S. Columbien, S. Am. Coll. Dr. Kriiger. W. Niepelt, Zirlau. / 
Brit. Mus. 1928-15 1. 1 o", Sierra Nevada de Sta. Marta, Columbien, S. Am. 1500 m. Juni 
1925. Coll. Dr. Kriiger. W. Niepelt, Zirlau. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 3, 
fig. 27.] 

Currently placed as a form of Eueides procula edias Hewitson. 



208 P. R. ACKERY & R. L. SMILES 

unifasciatus Butler, 1873a : 169 (as sp. of Eueides). 2 o* syntypes. Ega. Amazons. Bates 
57-20. [PI. 4, fig. 42.] 

Currently placed as a subspecies of Eueides vibilia (Godart). 

unimaculata Hewitson, 18696 : 10 (as sp. of Heliconia). 1 q* syntype. Ecuador. Hewitson 
Coll. 79.69. [PI. 20, fig. 200.] 

Currently placed as a transitional form between Heliconius melpomene plesseni Riffarth 
and Heliconius melpomene aglaope Felder & Felder. 

unipuncta Joicey & Kaye, 191 76 : 93, pi. 6, fig. 8 (as ab. of Heliconius erato amphitrite). 2 <J 
syntypes. Tirapata, Perou. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 33, fig. 338.] 
Currently placed as a form of Heliconius erato amphitvite Riffarth. 

vegetissima Stichel, 1903 : 8 (as ab. of Eueides isabella). Holotype $. Santa Jnez (Ecuad.). 
R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 5, 

fig- 53-] 

Currently placed as a form of Eueides isabella dissoluta Stichel. 

venustus Salvin, 1871 : 413 (as sp. of Heliconius). 1 o* syntype. Apolobamba, Bolivia. 
Pearce. / Godman-Salvin Coll. 1913-2. [PI. 33, fig. 339.] 
Currently placed as a subspecies of Heliconius erato (L.). 

veraepacis Bates, 1864a : 57 (as sp. of Heliconius). 4^1? syntypes. Forests of N. Vera 
Paz. F. D. G. & O. S. / Godman-Salvin Coll. 1913-12. [PI. 36, fig. 374.] 
Currently placed as a subspecies of Heliconius sara (Fabricius). 

vereatta Stichel, 1912 : 1 (as form of Heliconius hermathena). Holotype o*. Faro. v. 191 1. 
Ducke. / N. Brasilien, Faro. e. c. H. Stichel. / Rothschild Bequest. B. M. 1939-1. [PI. 28, 
fig. 288.] 

Currently placed as a subspecies of Heliconius hermathena Hewitson. 

vetustus Butler, 1873a : 165 (as sp. of Heliconius). 1 $ syntype. Demerara. Ex. Mus. Milne. 
1464b. [PI. 16, fig. 162.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius) . 

vialis Stichel, 1903 : 20, pi. 1, figs 14, 15 (as subsp. of Eueides vibilia). 1 $ syntype. Columb. / 
Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 4, fig. 39.] 

vicinalis Stichel, 1903 : 21 (as subsp. of Eueides vibilia). 2 $ syntypes. Palmar (Ecuad.). 
R. Haensch S. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 4, fig. 40.] 

viculata Riffarth, 1900 : 211 (as var. of Heliconius phyllis). 11 (J, 3 $ syntypes. Original. / 
Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. 1 <J, Cayenne. 1 o\ Chiriqui. 
1 <£, Paricatuba by Santarem, 14.6.93. 1 $, Venezuela, Porto Cabello. 3 q\ Surinam. 1 <J, 
Surinam, H. Stichel. 2 o". British Guayana. 1 q\ Bersaba, Surinam. 1898-9. Michls. 1 ?, 
Paricatuba. 13.6.93. 1 $, Santarem. 6.5.93. 1 $, Obidos. [PI. 31, fig. 318.] 
Currently placed as a form of Heliconius erato hydara (Hewitson) . 

virgata Stichel, 1902 : 355 (as ab. of Heliconius melpomene timareta). 1 <$ syntype. Santa 
Jnez, (Ecuad.). R. Haensch S. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 
1934-120. [PI. 38, fig. 392.] 

Currently placed as a form of Heliconius timareta Hewitson. 

viridana Stichel, 1906a : 35 (as form of Heliconius doris viridis). Holotype $. Colombia, 
Magdalena s. e. c. H. Stichel. / Typus. / Rothschild Bequest. B. M. 1939-1. [PI. 9, fig- 95-] 
Currently placed as a form of Heliconius doris eratonius Staudinger. 

vittatus Butler, 1873a : 166 (as sp. of Heliconius) . 1 5" syntype. Bogota. Stevens. 56-142. 
[PI. 15, fig. 1 54-] 

Currently placed as a form of Heliconius hecale ithaca Felder & Felder. 



TYPE-SPECIMENS OF HELICONIINAE 209 

vulcanus Butler, 1865 : 433, pi. 25, fig. 5 (as sp. of Heliconia). 1 q* syntype. 1763a. [PI. 18, 
fig. 180.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 
Described from Demerera and Panama. 

vulgiformis Butler & Druce, 1872 : 102 (as sp. of Eueides). 1 ? syntype. Costa Rica. Van 
Patten. Druce Coll. / Godman-Salvin Coll. 1913-2. [PI. 3, fig. 28.] 
Currently placed as a subspecies of Eueides procula Doubleday. 

warneri Hall, 1936 : 276 (as subsp. of Colaenis julia). 1 $ syntype. St. Kitts, B. W. I. 
December 1935, A. Hall. / Brit. Mus. 1936-736. [PI. 3, fig. 24.] 
Currently placed as a synonym of Dry as iulia iulia (Fabricius). 

wernickei Weymer, 1906 : 68 (as sp. of Heliconius). Holotype g. Bogota, Columbien. / Ex. 
Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 27, fig. 277.] 

Currently placed as a hybrid between Heliconius melpomene melpomene (L.) and Heliconius 
cydno cydno Doubleday. 

weymeri Staudinger, 1896 : 287 (as sp. of Heliconius) . 1 q\ 1 $ syntypes. R. Dagua, 
Columb. / Ex. Grose Smith. 1910. / Joicey Bequest. Brit. Mus. 1934-120. [PI. 27, fig. 273.] 
Currently placed as a subspecies of Heliconius cydno Doubleday. 

xanthicus Bates, 1864a : 57 (as sp. of Heliconius) . 1 $ syntype. Lion Hill, Panama. 
McLeannan. / Godman-Salvin Coll. 1913-2. [PI. 14, fig. 147.] 

Currently placed as a transitional form between Heliconius hecale melicerta Bates and 
Heliconius hecala zuleika Hewitson. 

xanthocles Bates, 1862 : 561 (as sp. of Heliconius). 1 q* syntype. Demerara, Brit. Guiana. 
Bates Coll. / Godman-Salvin Coll. 1913-2. [PI. 7, fig. 79.] 

xenoclea Hewitson, [1853] : pi. 1, fig. 1 (as sp. of Heliconia). 1 <J syntype. Hewitson Coll. 
79.69. [PI. 18, fig. 187.] 

Currently placed as a subspecies of Heliconius melpomene (L.). 

xenophanes Felder & Felder, 1865 : 377, pi. 46, figs 14, 15 (as sp. of Eueides). 1 o* syntype. 
Bogota. Lindig. Type. / Felder Colin. / Rothschild Bequest. B. M. 1939-1. [PI. 6, fig. 59.] 
Currently placed as a subspecies of Eueides tales (Cramer). 

zelinde Butler, 1869 : 17, pi. 9, fig. 1 (as sp. of Heliconius). 1 5* syntype. W. Coast of 
America. Kellct & Wood. 50-12. [PI. 27, fig. 270.] 

Currently placed as a subspecies of Heliconius cydno Doubleday. 

zobeide Butler, 1869 : 18, pi. fig. 3 (as sp. of Heliconius). 1 o* syntype. Para. Mrs Smith. 
45-70. [PI. 37, fig. 382.] 

Currently placed as a form of Heliconius antiochus alba Riffarth. 

zobrysi Frushstorfer, 1910 : 194 (as subsp. of Heliconius aulicus). 1 <? syntype. Brasilien, M. 
Grosso. Fruhstorfer. / Fruhstorfer Coll. B. M. 1937-285. [PI. 12, fig. 130.] 
Currently placed as a subspecies of Heliconius numata (Cramer). 

zuleika Hewitson, [1853] : pi. 3, fig. 10 (as sp. of Heliconia). 2 5*, 1 $ syntypes. Nicaragua. 
Hewitson Coll. 79.69. [PI. 13, fig. 140.] 

Currently placed as a subspecies of Heliconius hecale (Fabricius) . 

zygia Riffarth, 19076 : 504, pi. 5, fig. 5 (as form of Heliconius andevida melicerta) . 1 o" syntype. 
Columb. / Original. / Ex. Coll. Riffarth. / Joicey Bequest. Brit. Mus. 1934-120. [PL 14, 
fig. 144] 

Currently placed as a form of Heliconius hecale melicerta Bates. 



P. R. ACKERY & R. L. SMILES 



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■ 1865. Reise der Osterreichischen Fregalte Novara; Lepidoptera, Rhopalocera 

[published 1864-1867] vi + 548 pp., 140 pis. Wien. 
Fruhstorfer, H. 1910. Drei neue Heliconidenformen. Ent. Z., Frank/, a. M. 24 : 194. 

191 2. Neue Nymphaliden des neotropischen Gebiets aus der Sammlung Staudinger. 

Ent. Rdsch. 29 : 14-15. 

Godman, F. D. & Salvin, O. 1877. Descriptions of twelve new species and a new genus of 

Rhopalocera from central America. Proc. zool. Soc. Lond. 1877 : 60-64. 
— ■ 1 88 1. Biologia Centrali- Americana, Insecta, Lepidoptera-Rhopalocera 1 : xlv -f- 

487 pp. [Published 1879-1901]. London. 
Grose-Smith, H. 1898. Descriptions of new species of butterflies from South America. 

Ann. Mag. nat. Hist. (7) 2 : 70-73. 
& Kirby, W. F. 1892. Nymphalidae-Heliconinae, Heliconius, 1 pi. In Rhopalocera 

Exotica 1 : parts 1-20 [published 1 887-1 892]. London. 
Hall, A. 1921. Descriptions of three new butterflies from Colombia. Entomologist 54 : 

278-279. 
— — 1930. New forms of tropical Rhopalocera. Entomologist 63 : 278-279. 

■ 1936. The butterflies of St. Kitts. Entomologist 69 : 274-278. 

Hering, M. 1926. Zur Kenntnis der stidamerikanischen Nymphaliden-Gattung Dione Hb. 

Dt. ent. Z. 40 : 195-204, 5 figs. 
Hewitson, W. C. [1853] Heliconidae, Heliconia, 3 pis. In Illustrations of new species of 

exotic butterflies. 1 : parts 1-20 [published 1851-1856]. London. 

1858. Heliconidae, Tithorea & Heliconia, 1 pi. In Illustrations of new species of exotic 

butterflies 2 : parts 21-40 [published 1857-1861]. London. 

1861. Descriptions of new diurnal lepidoptera. /. ent. London 1 : 155-158, 2 pis. 

1864. Descriptions of new species of diurnal Lepidoptera. Trans, ent. Soc. Lond. (3) 

2 : 245-249, 2 pis. 

1867a. Heliconidae, Heliconia, 2 pis. In Illustrations of new species of exotic butterflies 

4 : parts 61-79 [published 1867-1871]. London. 

18676. Descriptions of some new species of diurnal Lepidoptera. Trans, ent. Soc. 

Lond. (3) 5 : 561-566. 

1869a. Descriptions of six new species of diurnal Lepidoptera from Nicaragua. Trans. 

ent. Soc. Lond. 1869 : 33-35. 

18696. Equatorial Lepidoptera Parts I & II: ii + 32 pp. London. [Reprinted in 

Hewitson on Butterflies, 1876-1877, published by Classey, Hampton, 1972.] 

1872a. Descriptions of seven new exotic Rhopalocera. Entomologist's mon. Mag. 

9 : 83-86. 

18726-1873. Heliconidae, Tithorea & Heliconia, Heliconia, 3 pis. In Illustrations of new 

species of exotic butterflies 5 : parts 81-100 [published 1872-1877]. London. 



212 P. R. ACKERY & R. L. SMILES 

1874. Description of a new species of Heliconia from central America. Entomologist's 

mon. Mag. 10 : 224. 

1875. Descriptions of six new species of butterflies from South America, West Africa 

and Borneo. Entomologist's mon. Mag. ±1 : 182-184 

Holzinger, H. & Holzinger, R. 1974- Die Typen in der Heliconiinae-Sammlung des 

Naturhistorischen Museums in Wien (Lep., Nymphalidae). Annln naturh. Mus. Wien 

78 : 261-273, 1 pi. 
Joicey, J. J. & Kaye, W. J. 1917a. On a collection of Heliconine forms from French Guiana. 

Trans, ent. Soc. Lond. [1916] : 412-431, 2 pis. 
19176. New races and aberrations of Heliconius. Ann. Mag. nat. Hist. (8) 20 : 

87-94, 2 pis. 

1919. Notes on a large Heliconine collection made in French Guiana in 1917, 



compared with a similar collection made in 1915. Trans, ent. Soc. Lond. [1918] : 347-353. 
— • — ■ & Talbot, G. 1925. Notes on some Lepidoptera with descriptions of new forms. 

Ann. Mag. nat. Hist. (9) 16 : 633-653. 
Kaye, W. J. 1916. New forms of Heliconius. Entomologist's Rec. J. Var. 28 : 194-195. 

1919. New South American butterflies. Ann. Mag. nat. Hist. (9) 3 : 215-218. 

Kruger, E. G. 1925. Einige neue Falterformen aus Kolumbien und Surinam. Dt. ent. Z. 

Iris 39 : 146-151. 
Kruger, R. 1927. Eine neue Heliconius -form. Int. ent. Z. 21 : 86. 
— ■ — - 1929. Neubeschreibungen. Int. ent. Z. 22 : 376-377. 
Lathy, P. I. 1906. On three new forms of butterfly of the genus Heliconius. Proc. zool. 

Soc. Lond. 1906 : 452-453, 1 pi. 
Michener, C. D. 1942. A review of the subspecies of Agraulis vamllae (Linnaeus). 

Lepidoptera: Nymphalidae. Am. Mus. Novit. No. 1215 : 1-7. 
Neustetter, H. 1925. Drei neue Heliconius-Formen. Int. ent. Z. 19 : 60-61. 
— ■ — 1926a. Beschreibung und Besprechung neuer und wenig gekannter Heliconius-Formen. 

Int. ent. Z. 20 : 34-40. 

19266. Besprechung einer Heliconius-Ausbeute aus Franzosisch-Guayana und 

Beschreibung der neuen Formen aus derselben. Int. ent. Z. 20 : 278-282, 289-293. 

1927a. Eine neue Heliconius-Form. Int. ent. Z. 21 : 86. 

19276. Neue Heliconius-Formen. Int. ent. Z. 21 : 227-230. 

1928a. Zwei neue Heliconius. Int. ent. Z. 21 : 442-444. 

19286. Neue Heliconius-Formen. Int. ent. Z. 22 : 75-80. 

1928c. Neue und wenig bekannte Heliconius. Int. ent. Z. 22 : 245-248. 

19280". Neue Heliconius. Int. ent. Z. 22 : 258-259. 

Niepelt, W. 1907a. Zwei neue Formen der Gattung Heliconius. Berl. ent. Z. 52 : 213-215. 

19076. Zwei neue Lokalformen der Gattung Heliconius Latr. Societas ent. 22 : 42. 

1908. Neue Formen der Gattung Heliconius (Lep.). Dt. ent. Z. 1908 : 505-506. 

1909. Neue Heliconius-Formen. Berl. ent. Z. 54 : 106-107. 

1914- In Strand, E., Lepidoptera Niepeltiana 1 : 64 pp., 12 pis. 

1915. Neue Formen exotische Tagfalter. Int. ent. Z. 9 : 58. 

1921. Eine neue Heliconius Form v. Guatemala. Int. ent. Z. 15 : 19. 

1925. Eine neue Heliconius-Form von Bolivia. Int. ent. Z. 18 : 252. 

1926a. Nachtrag und Berichtigung zu: Neue und wenig bekannte sudamerikanische 

Tagfalter. Int. ent. Z. 20 : 137-138. 

19266. Eine neue Heliconius-Form von Bolivia. Int. ent. Z. 19 : 337. 

Oberthur, C. 1902. Variation des Heliconia thelxiope et vesta. Etudes d' Entomologie 21 : 

1-26, 12 pis. 

■ 1916. Explication des planches. Etude lepidopterologie comparee 12 (2) : 44 pp., 43 pis. 

■ 1920. Explications des planches coloriees. £tude lepidopterologie comparee 17 : 59 pp., 

28 pis. 
1923. Explication des planches (DLXII-DLXIV). £tude lepidoptirologie comparie 20 : 

301-304, 3 pis. 



TYPE-SPECIMENS OF HELICONIINAE 213 

1925. Lepidopteres de l'Amerique du Sud. Etude lepidopterologie compare'e 22 : 81-89, 

9 figs- 
Poulton, E. B. 1910. In Joseph, E. G., The collections of William John Burchell D. C. L. 

in the Hope Department, Oxford University Museum. On the Lepidoptera, Rhopalocera 

collected by W. J. Burchell in Brazil, 1825-1830. Ann. Mag. nat. Hist. (8) 5 : 322-346. 
Riffarth, H. 1899. Neu Heliconius-F or men. Berl. ent. Z. 43 : 405-408. 
■ 1900. Die Gattung Heliconius Latr. Neu bearbeitet und Beschreibung neuer Formen. 

Berl. ent. Z. 45 : 183-214. 
— — ■ 1901. Die Gattung Heliconius Latr. Neu bearbeitrr und Beschreibung neuer Formen. 

Berl. ent. Z. 46 : 25-185. 

1902. Nochmals Ch. Oberthiir's Etudes d'Entomologie, vol. 21. Berl. ent. Z. 47 : 157-166 

' 1906. Eine neue Heliconius-Form. Insektenborse 23 : 56. 

1907a. Eine neue Lokalform von Heliconius batesi Riff. (Lep.). Dt. ent. Z. 1907 : 

333-334- 

19076. Neue und wenig bekannte Formen der Gattung Heliconius, nebst einer neuen 

Eueides-Form. (Lep.). Dt. ent. Z. 1907 : 501-514, 1 pi. 

1908. Uber Neustetters neue Heliconins-Formen (Lepid.). Dt. ent. Z. 1908 : 110-114. 

Riley, N. D. 1919. Some new Rhopalocera from Brazil collected by E. H. W. YVickham, 

Esq. Entomologist 52 : 181-186. 
1926. Colaenis and Dione (Lep. Nymphalidae) : A revisional note on the species. Ento- 
mologist 59 : 240-245, 1 pi. 
Rober, J. 1927. Neue exotische Falter. Int. ent. Z. 20 : 400-403. 
Salvin, O. 1 87 1. Descriptions of new species of butterflies from tropical America. Ann. 

Mag. nat. Hist. (4) 7 : 412-416. 
Salvin. O. & Godman, F. D. 1868. On some new species of diurnal Lepidoptera from South 

America. Ann. Mag. nat. Hist. (4) 2 : 141-152. 
Schaus, W. 1921. New species of Lepidoptera in the United States National Museum. 

Proc. U. S. natn. Mus. 57 : 107-152. 
Seitz, A. 19 i 3. Subfamily Heliconiinae. In Seitz, A., Die Gross-Schmetterlinge der Erde 

Exotische Fauna 5 : 375-402, 10 pis. 
Srnka, A. 1885. Neue siidamerikanische Danaidae und Heliiconidae. Berl. ent. Z. 29 : 

121-130, 1 pi. 
Staudinger, O. 1896. Neue Heliconius-Arten und Formen. Dt. ent. Z. Iris 9 : 284-317, 

2 pis. 
Stichel, H. 1899a. Neue Heliconius aus Siid-Brasilien. Ent. Nachr. 25 : 28-30. 

18996. Heliconius nanna n. sp. Int. ent. Z. 12 : 143. 

1902. Nachrichten aus dem Berliner Entomologischen Verein. Insektenborse 19 : 355. 

1903. Synonymisches Verzeichnis bekannter Eueides-Formen mit erlauternden Bemer- 

kungen und Neubeschreibungen. Berl. ent. Z. 48 : 1-34, 1 pi. 

1906a. Lepidoptera Rhopalocera, Fam. Nymphalidae Subfam. Heliconiinae. In 

Wytsman, P., Genera Insect. 37 : 74 pp., 6 pis. 

19066. In Nachrichten aus den Berliner Entomologischen Verein. Insektenborse 

23 : 208. 

1907. Lepidoptera Rhopalocera, Fam. Nymphalidae Subfam. Dioninae. In Wytsman, 

P., Genera Insect. 63 : 38 pp., 3 pis. 

1909. Beschreibung einiger neue Heliconiidae. Societas ent. 23 : 178-179. 

191 2. Zwei neue Formen von Heliconius hermathena Hew. Int. ent. Z. 4 : 1-2. 

1919- Anmerkungen und Zutrage zur Gattung Heliconius L. Neue Beitr. syst. Insektenk. 

1 : 119-120. 

1923. Kolombische Heliconius (Lep., Rhop.). Dt. ent. Z. 1923 : 260-270. 

■ & Riffarth, H. 1905. Heliconiidae. Tierreich 22 : xvi + 290 pp. 

Stoll, C. [1 780-1 782]. Die Uitlandsche Kapellen ... 4 [part] : 29-252, pis. 305-400 [com- 
pletion of work started by Cramer, P.]. Amsteldam & Utrecht. 



2i 4 P- R- ACKERY & R. L. SMILES 

Talbot, G. 1928. List of Rhopalocera collected by Mr. C. L. Collenette in Matto Grosso, 
Brazil. Bull. Hill. Mus. Witley 2 : 192-220. 

1932. New species and forms of Lepidoptera from South America. Bull. Hill. Mus. 

Witley 4 : 189-197. 

Turner, J. R. G. 1966a. A little-recognised species of Heliconius butterfly (Nymphalidae) . 
J. Res. Lepid. 5 : 97-112, 1 pi. 

19666. A rare mimetic Heliconius (Lepidoptera: Nymphalidae). Proc. R. ent. Soc. Lond. 

(B) 35 : 128-132. 

1971- Two thousand generations of hybridisation in a Heliconius butterfly. Evolution, 

Lancaster, Pa. 25 : 471-482. 

1973- Illustrations of Heliconius; some rare and important species. /. Lepid. Soc. 

27 : 130-136. 

Vane-Wright, R. I. 1975. The butterflies named by J. F. Gmelin (Lepidoptera: Rhopalo- 
cera). Bull. Br. Mus. nat. Hist. (Ent.) 32 : 17-64, pis 1-6. 

Ackery, P. R. & Smiles, R. L. 1975. The distribution, polymorphism and mimicry of 

Heliconius telesiphe (Doubleday) and the species of Podotricha Michener (Lepidoptera: 
Heliconiinae) . Trans. R. ent. Soc. Lond. 126 : 611-636, 1 pi., 6 figs. 

Weymer, G. 1906. Zwei neue Heliconius-Formen. Dt. ent. Z. Iris 19 : 68-71. 

1912. Einige neue Heliconiden-Formen. Ent. Rdsch. 29 : 73-74. 



P. R. Ackery 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 

R. L. Smiles 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 



D 



PLATE i 



Fig. 


i. 


Fig. 


2. 


Fig. 


3- 


Fig. 


4- 


Fig. 


5- 


Fig. 


6. 


Fig. 


7- 


Fig. 


8 


Fig. 


9 


Fig. 


IO 



Upper- and underside 

Metamandana dido pygmalion Fruhstorfer, holotype 9 (Obidos). 
M. dido diatonica Fruhstorfer, holotype g (Honduras, San Pedro Sula). 
Colaenis phaetusa form stupenda Stichel, holotype <$ (Bugaba, Panama). 
Colaenis phaetusa form deleta Stichel, holotype £ (Paraguay, Nuev. Germania). 

C. phaetusa form lutulenta Stichel, holotype ^ (Paraguay, Nuev. Germania). 
Dione miraculosa Hering, paratype <§ (Arequipa, Stid-Peru). 

Agraulis andicola Bates, syntype <J (Tacunga, Ecuador). 

Dione glycera gnophota Stichel, syntype $ (Colombia, Cordill. p. Bogota). 

D. moneta butleri Stichel, syntype g (Colombia, R. Magdalena). 

D. vanillae incavnata Riley, lectotype £ (Near Durango City, Mexico). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 1 




PLATE 2 



Fig. 


ii. 


Fig. 


12. 


Fig. 


13 


Fig. 


14 


Fig. 


15 


Fig. 


16 


Fig. 


17 


Fig. 


18 


Fig. 


19 


Fig. 


20 


Fig. 


21 


Fig. 


22. 



Upper- and underside 

Dione vanillae maculosa Stichel, syntype £ (Brasilia, Espirito Santo). 

Agraulis lucina Felder & Felder, syntype $ (Rio Negro). 

Dione vanillae catella Stichel, holotype $ (Siid-Peru, Pozuzo). 

Colaenis euchroia Doubleday, lectotype <$ (Columbia). 

C. euchroia caucana Riley, lectotype <£ (Frontino, Antioquia). 

C. euchroia mellosa Stichel, lectotype £ (Ecuador). 

C. euchroia straminea Riley, holotype q (Balzapamba, Ecuador). 

C. tithraustes Salvin, lectotype $ (Rio Topo, Ecuador). 

C. telesiphe Hewitson, lectotype $ (Ecuador). 

C. tithraustes var. diaphana Niepelt, holotype <J (S. O. Peru). 

C. julia carteri Riley, syntype <J (Nassau, Bahamas). 

C. julia lucia Riley, syntype g (Santa Lucia). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 2 







1^1 



/ 








I" 









i5^*y 









^V 




PLATE 3 

Upper- and underside 

Fig. 23. Colaenis julia framptoni Riley, syntype <§ (W. Indies, St. Vincent). 

Fig. 24. C. julia wavneri Hall, syntype 9 (B.W.I., St. Kitts). 

Fig. 25. C. iulia titio Stichel, syntype <$ (Sta. Cruz, Slid Bolivia). 

Fig. 26. Eueides edias Hewitson, syntype <$ (New Granada). 

Fig. 27. E. edias umbratilis Rober, syntype £ (Sierra Nevada de Sta. Marta, Columbien). 

Fig. 28. E. vulgiformis Butler & Druce, syntype 9 (Costa Rica). 

Fig. 29. E. eurysaces Hewitson, syntype 9 (Quito). 

Fig. 30. E. procula Doubleday, syntype $ (Venezuela). 

Fig. 31. E. lampeto copiosus Stichel, syntype 9 (British Guiana, Potaro Rd.). 

Fig. 32. E. lampeto Bates, holotype 9 (S. Paulo). 

Fig. 33. E. lampeto ab. fuliginosa Stichel, syntype g (Ecuador, Santa Jnez). 

Fig. 34. E. acacetes Hewitson, syntype $ (Ecuador). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 3 








\- — 
24%s» 




















^33 






;:^34 



Fig. 


35- 


Fig. 


36. 


Fig. 


37- 


Fig. 


38- 


Fig. 


39- 


Fig. 


40. 


Fig. 


41. 


Fig. 


42. 


Fig. 


43- 


Fig. 


44. 


Fig. 


45- 


Fig. 


46. 



PLATE 4 

Upper- and underside 

Eueides lampeto ab. amoena Stichel, syntype <§ (Ecuador, Santa Jnez). 

E. lampeto acacetes ab. carbo Stichel, syntype 9 (Ecuador, Santa Jnez). 

E. lampeto fuliginosus form pallida Riffarth, holotype £ (Ecuador, Ob. Pastaza). 

E. emsleyi Brown, holotype <£ (Colombia, Valdevia). 

E. vibilia vialis Stichel, syntype g (Columbia). 

E. vibilia vicinalis Stichel, syntype 9 (Ecuador, Palmar). 

E. vibilia vibilia form pallens Stichel, holotype § (Brazil, Espirito-Santo). 

E. unifasciatus Butler, syntype <J (Ega, Amazons). 

E. thy ana Felder & Felder, syntype § (Brazil). 

E. lineata Salvin & Godman, syntype £ (Polochic Valley). 

E. eanes Hewitson, syntype £ (Amazons). 

E. eanes ab. riffarthi Stichel, syntype $ (Siid-Peru, Chanchamayo). 



Bull. Br. Mus. licit. Hist. (Ent.) 32, 5 



PLATE 4 







**f^ r 










39^ 




\4^\> 













44 







46 



PLATE 5 

Upper- and underside 

Fig. 47. Eueides eanes ab. eanides Stichel, syntype £ (Bolivien). 

Fig. 48. E. cleobaea zorcaon ab. adnsta Stichel, syntype $ (Mexico, Vera Cruz). 

Fig. 49. E. pellucida Srnka, holotype 9- 

Fig. 50. E. Isabella arquata Stichel, syntype $ (S. America, Caucathal). 

Fig. 51. Heliconius Isabella huebneri ab. spoliata Stichel, holotype $ (S. America, 

Caucathal). 

Fig. 52. Eueides Isabella seitzi Stichel, syntype £ (Santa Jnez, Ecuad.). 

Fig. 53. E. Isabella ab. vegetissima Stichel, holotype ^ (Santa Jnez, Ecuad.). 

Fig. 54. E. hippolinus Butler, syntype 9 (E. Peru). 

Fig. 55. E. Isabella hippolina ab. brunnea Stichel, syntype ^ (Tarapota, Peru). 

Fig. 56. E. Isabella dissoluta Stichel, syntype^ (Tarapota, Peru). 

Fig. 57. E. dynastes Felder & Felder, syntype $ (Venezuela). 

Fig. 58. Heliconius thales surdus Stichel, syntype <§ (Demerara). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 5 














\ /* 




56 









PLATE 6 



Fig. 


59 


Fig. 


60 


Fig. 


61 


Fig. 


62 


Fig. 


63 


Fig. 


64. 


Fig. 


65 


Fig. 


66. 


Fig. 


67. 


Fig. 


68. 


Fig. 


69 


Fig. 


70. 



Upper- and underside 

Eueides xenophanes Felder & Felder, syntype <§ (Bogota). 

E. thales heraldicus Stichel, syntype 9 (Para). 

E. cvystalina Hall, syntype g (Crystalina, W. Colombia). 

E. tales calathus Stichel, syntype ^ (Ob. Pastaza, Ecuad.). 

E. aliphera gracilis Stichel, syntype £ (Costa Rica, San Carlos). 

E. leucomma Bates, syntype g (Lion Hill, Panama). 

E. heliconioides Felder & Felder, syntype g (Ecuador). 

Heliconia emmelina Oberthiir, holotype g (Guyane anglaise). 

Heliconius bartletti Druce, syntype £ (Santa Cruz, Peru). 

H. aoede cupidineus Stichel, syntype <§ (Tarapoto, Peru). 

H. aoede eurycleia Brown, paratype £ (Cuiaba-Porto Velho). 

Heliconia thetis Boisduval, holotype 9 (Nicaragua). 



Bull. Br. Mus. nal. Hist. (Ent.) 32, 5 



PLATE 6 












63%^ 



fed 





PLATE 7 

Upper- and underside 

Fig. 71. Heliconia hierax Hewitson, syntype $ (Ecuador). 

Fig. 72. H. hecuba Hewitson, syntype $ (N. Granada). 

Fig. 73. Heliconius tolima Fassl, syntype $ (Canon del Tolima, Columb.). 

Fig. 74. H. cassandra Felder & Felder, syntype $ (Bogota). 

Fig. 75. H. hecuba intermedins Riffarth, holotype £ (Columbien). 

Fig. 76. H. crispus var. crespinus Krliger, syntype 9 (Cauca V.). 

Fig. 77. Heliconia choarina Hewitson, syntype £ (Ecuador). 

Fig. 78. Heliconius hecuba choarinus form flava Neustetter, holotype $ (Baiza, Ecuad. 

Fig. 79. H. xanthocles Bates, syntype <$ (Demerara, Brit. Guiana). 

Fig. 80. Heliconia catemaulti Oberthur, syntype^ (Cayenne). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 7 




PLATE 8 



Upper- and underside 

Fig. 8i. Heliconius pavaplesius Bates, syntype £ (Maranham, N. Brazil). 

Fig. 82. Heliconia cethosia form completa Oberthiir, syntype £ (Region de Bogota). 

Fig. 83. Heliconius melete Felder & Felder, syntype £ (Nova Granada). 

Fig. 84. H. mimulinus Butler, syntype (J (Bogota). 

Fig. 85. H. clytia var. elsa Riffarth, syntype $ (Surinam). 

Fig. 86. H. wallacei araguaia Brown, holotype <$ (R. Araguaya, Prov. Goyaz). 

Fig. 87. H. clytia ab. parvimaculata Riffarth, syntype 9 (Obidos, Amazonen). 

Fig. 88. H. wallacei brevimaculata ab. halli Kaye, holotype 9 (Serpa, Lower Amazon). 

Fig. 89. H. lindigii Felder & Felder, syntype <$ (Bogota). 

Fig. 90. H. burneyi serpensis Kaye, paratype £ (Serpa, Lower Amazon). 

Fig. 91. H. egeria ab. clearista Oberthiir, syntype $ (Guyane francaise). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 8 














89 






90 



3J|w 





91 



PLATE 9 



i-IG. 


92. 


Fig. 


93- 


Fig. 


94- 


Fig. 


95- 


Fig. 


96. 


Fig. 


97- 


Fig. 


98. 


Fig. 


99. 


Fig. 


100 


Fig. 


101. 



Upper- and underside 

Heliconius astraea rondonia Brown, paratype £ (Matto-Grosso, Brazil). 

H. erato luminosus Riffarth, syntype £ (Columb.). 

H. erato ab. tecta Riffarth, syntype <§ (Columbien). 

H. doris viridis form viridana Stichel, holotype $ (Colombia, Magdalena). 

H. erato aristomache Riffarth, syntype^ (Balzapamba). 

H. doris doris ab. gibbsi Kaye, holotype <$ (Friendship, B. Guiana). 

H. doris caernleatus Stichel, syntype g (Slid Peru, Pozuzo). 

H. nattereri Felder & Felder, syntype g (Bahia, Beske). 

H . fruhstorferi Riffarth, holotype 9 (Espirito-Santo, Brazil). 

H. messene Felder & Felder, syntype <§ (Bogota). 



Bull. Br. Mus. nal. Hist. (Ent.) 32, 5 



PLATE 9 





Fig. 


102. 


Fig. 


I03. 


Fig. 


IO4. 


Fig. 


IO5. 


Fig. 


I06. 


Fig. 


I07. 


Fig. 


I08. 


Fig. 


IO9. 


Fig. 


110. 


Fig. 


III. 


Fig. 


112. 



PLATE 10 

Upper- and underside 

Heliconius euphone Felder & Felder, paralectotype g (Bogota). 

H. aristiona lepidus Riffarth, syntype § (Sara-yacu, Ecuad.). 

H. aristiona lepidus form gracilis Riffarth, syntype $ (La Merced, Peru). 

H. aristiona tarapotensis Riffarth, syntype £ (Tarapota, Peru). 

Heliconia aristiona Hewitson, syntype <§ (Colombia) . 

Heliconius aristiona pratti Joicey & Kaye, syntype 9 (Charape, N. Peru). 

H. bicoloratus Butler, syntype <§ (Nauta, Amazons). 

H. numatus praelautus Stichel, holotype 9 (Ucayali). 

Heliconia arcuella Druce, syntype <$ (Nauta, Ecuador). 

Heliconius aurora Bates, syntype ^ (S. Paulo, Amazons). 

H. nubifer Butler, holotype 9 (Fonteboa, Amazons). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 10 



r- 





* 



102 



19 m 



s 





107 




PLATE ii 

Upper- and underside 

Heliconius numata var. isabellinus Bates, syntype $ (S. Paulo, U. Amazons). 
H. ignotus Joicey & Kaye, syntype <J (Charape, N. Peru). 

H. num ata talboti Joicey & Kaye, syntype <$ (Rentema Falls, Upper Maranon) 
H. novatus subnubilis Stichel, syntype § (Juanjuy, Peru). 
H. silvana mirificus Stichel, holotype 9 (Ucayali). 
H. novatus artemis Riffarth, holotype,^ (Titicaca, S. Bolivia). 
H. novatus obscurior Stichel, syntype £ (Bolivia, Titicaca). 
H. peeblesi Joicey & Talbot, syntype g (Merida Dist., Venezuela). 
H. numata superioris ab. translata Joicey & Kaye, syntype 9 (Manaos, Lower 
Amazon). 
Fig. 122. Heliconius numata ab. or var. guiensis Riffarth, holotype 9 (British Guayana). 



tig. 


"3- 


Fig. 


114. 


Fig. 


"5- 


Fig. 


116. 


Fig. 


117. 


Fig. 


118. 


Fig. 


119. 


Fig. 


120. 


Fig. 


121. 



Bull. Br. Mus. nul. Hist. (Ent.) 52., 5 



PLATE 1 [ 




Fig. 


123. 


Fig. 


124. 


Fig. 


125. 


Fig. 


126. 


Fig. 


127. 


Fig. 


128. 


Fig. 


129. 


Fig. 


130. 


Fig. 


131- 



PLATE 12 

Upper- and underside 

Heliconius seraphini Talbot, holotype g (Guyane, St. Laurent). 

H. numata form melanopors Joicey & Kaye, syntype 9 (Guyane Francse, St. Jean 

du Maroni). 

H. numatus sincerus Riffarth, syntype <J (Amazon) . 

H. superioris Butler, syntype g (Villa Nova, Amazons). 

H. numata silvaniformis Joicey & Kaye, syntype <$ (Para, Lower Amazon). 

H. diffusus Butler, syntype $ (Para). 

Heliconia telchinia Doubleday, syntype 9 (Honduras). 

Heliconius aulicus zobrysi Fruhstorfer, syntype £ (Brasilien, M. Grosso). 

H. clarescens Butler, holotype 9 (Bugaba, Panama). 



Bull. Br. Mus. nut. His/. (Ent.) 32, 5 



PLATE 12 




PLATE 13 

Upper- and underside 

Fig. 132. Heliconius ismenius occidentalis Neustetter, syntype <J (Cantinella, Putamayo). 

Fig. 133. H. ismenius ismenius form immoderata Stichel, syntype $ (Columb., Muzo). 

Fig. 134. H. ismenius var. or ab. hermanni Riffarth, holotype $ (Columb.). 

Fig. 135. H. fasciatus Godman & Salvin, syntype <J (Lion Hill, Panama). 

Fig. 136. H. pardalinus butleri Brown, holotype $ (Perou, Cavallo-Cocho). 

Fig. 137. H. eucoma var. pardalinus Bates, syntype <$ (S. Paulo, U. Amazons). 

Fig. 138. H. anderida form styx Niepelt, syntype g. 

Fig. 139. Heliconia fomarina Hewitson, syntype 3 1 (S. America). 

Fig. 140. H. zuleika Hewitson, syntype $ (Nicaragua). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 13 




PLATE 14 

Upper- and underside 

Fig. 141. Heliconius chrysantis Godman & Salvin, holotype <$ (Nicaragua). 

Fig. 142. H. zuleika ab. albipanctata Riffarth, syntype $ (Chiriqui). 

Fig. 143. H. melicerta Bates, syntype <$ (Lion Hill, Panama). 

Fig. 144. H. anderida melicerta form zygia Riffarth, syntype £ (Columb.). 

Fig. 145. H. metaphovus var. semipkorus Staudinger, syntype g (Rio Dagua) 

Fig. 146. H. jucundus Bates, syntype <$ (Lion Hill, Panama). 

Fig. 147. H . xanthicus Bates, syntype $ (Lion Hill, Panama). 

Fig. 148. H. albucilla Bates, syntype $ (Lion Hill, Panama). 

Fig. 149. H. euchevius Weymer, holotype $ (Bogota, Columbien). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 14 




Fig. 


150. 


Fig. 


151 


Fig. 


152 


Fig. 


i53 


Fig. 


J 54 


Fig. 


155 


Fig. 


156 


Fig. 


157- 


Fig. 


158. 



PLATE 15 

Upper- and underside 

Heliconius hecale australis Brown, paratype £ (Quevedo, W. Ecuador) . 

Heliconia anderida Hevvitson, syntype q (Caracas). 

Heliconius ithaca Felder & Felder, syntype $ (Bogota). 

H. aristiona indecisa Joicey & Kaye, syntype g (Up. Orinoco). 

H. vittatus Butler, syntype <J (Bogota). 

Heliconia quitalena Hewitson, syntype <$ (Quito). 

Heliconius humboldti Neustetter, syntype $ (Rio Itaya bv Iquitos). 

H. humboldti form alexander Neustetter, syntype $ (Rio Itaya by Iquitos). 

H. sisyphus Salvin, syntype £> (Cosnipata Vail., E. Peru). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 15 




Fig. 


159- 


Fig. 


160. 


Fig. 


161. 


Fig. 


162. 


Fig. 


163. 


Fig. 


164. 


Fig. 


165. 


Fig. 


166. 


Fig. 


167. 


Fig. 


168. 



PLATE 16 

Upper- and underside 

Heliconius pasithoe var. fulvescens Lathy, holotype 9 (Demerara) 

H. hecale barcantl Brown, holotype £ (C irupano). 

H. hecale clearei Hall, syntype <$ (Mabaruma, British Guiana). 

H. vetustus Butler, syntype 9 (Demerara). 

H. novatus Bates, syntype 9 (Para). 

H. schulzi Riffarth, syntype 9 (Para). 

H. radiosus Butler, syntype 9 (Villa Nova). 

H. paraensis var. latus Riffarth, syntype g (Itaituba). 

H. paraensis Riffarth, syntype ^ (Para). 

H . clandia Godman & Salvin, syntype $ (Calobre, Panama). 



Bull. Br. Mus. not. Hist. (Ent.) 32, 5 



PLATE 16 




168 



PLATE 17 

Upper- and underside 

Fig. 169. Heliconius metalilis Butler, syntype g (Venezuela). 

Fig. 170. H. cephallenia Felder & Felder, syntype <$ (Surinam, Guiana). 

Fig. 171. H. ethilla chapadensis Brown, paratype <$ (Matto Grosso). 

Fig. 172. H. aerotome Felder & Felder, syntype <§ (Rio Negro). 

Fig. 173. Heliconia hippola Hewitson, syntype <$. 

Fig. 174. H. atthis Doubleday, syntype £ (Guayaquil). 

Fig. 175. Heliconius elevatus roraima Turner, holotype £ (Roraima, B. Guiana). 

Fig. 176. Heliconia bari Oberthiir, syntype $ (Cayenne). 

Fig. 177. Heliconius elevatus schmassmanni Joicey & Talbot, holotype <$ (Cuyaba-Corumba, 

Matto Grosso). 

Fig. 178. H. elevatus perchlora Joicey & Kaye, holotype <J (Boliv.). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE i' 





170 




1 





175 





177 



PLATE i 8 

Upper- and underside 

Fig. 179. Heliconia rosina Boisduval, syntype £ (Mexiq.). 

Fig. 180. H. vulcanus Butler, syntype £. 

Fig. 181. Heliconius vulcanus ab. modesta Riffarth, syntype £ (Paramba, Ecuador). 

Fig. 182. H. vulcanus sticheli Riffarth, syntype^ (Paramba, Ecuador). 

Fig. 183. Heliconia cythera Hewitson, syntype $ (Ecuador). 

Fig. 184. Heliconius vulcanus form concinna Stichel, holotype £ (Balzapamba, Ecuad. 

Fig. 185. H. xenoclea var. confluens Lathy, syntype <$ (Pichis Rd., Peru). 

Fig. 186. H. batesi Riffarth, syntype <§ (Peru?). 

Fig. 187. Heliconia xenoclea Hewitson, syntype £. 

Fig. 188. Heliconius xenoclea var. superba Lathy, holotype g (Rio Colorado, Peru). 



Bull. Br. Mns. mil. Hist. (Ent.) 32, 5 



X 



PLATE 1! 




179 








184 







183 




188 



PLATE 19 

Upper- and underside 

Fig. 189. Heliconius melpomene riffarthi Stichel, syntype $ (Ucayali). 

Fig. 190. H. melpomene riffarthi form rubescens Stichel, holotype <$ (Chuchurras, Peru). 

Fig. 191. H. amaryllis Felder & Felder, syntype £ (Rio Negro). 

Fig. 192. H. melpomene melpophylla Joicey & Kaye, holotype $. 

Fig. 193. H. batesi form plesseni Riffarth, syntype $ (Pastazza super., Ecuador). 

Fig. 194. H. batesi plesseni form rubicunda Niepelt, syntype $. 

Fig. 195. H. batesi plesseni form corona Niepelt, syntype J 1 . 

Fig. 196. H. batesi plesseni form pura Niepelt, syntype $ (Ob. Pastaza, Ecuad.). 

Fig. 197. H. batesi plesseni form adonis Riffarth, syntype $. 

Fig. 198. H. melpomene aglaope form fraterna Niepelt, syntype <$ (Ecuador, Canelos). 



Bull. Br. Mus. not. Hist. (Ent.) 32, 5 



PLATE 19 






190 




*W ^W 194 



*^P ^Pl95 





192 





197 



193^^^^ 




198 



Fig. 


199. 


Fig. 


200. 


Fig. 


201. 


Fig. 


202. 


Fig. 


203. 


Fig. 


204. 


Fig. 


205. 


Fig. 


206. 


Fig. 


207. 


Fig. 


208. 



PLATE 20 

Upper- and underside 

Heliconius xenoclea plesseni form clytie Neustettcr, holotype £ (Guayes, Ecuador) 

Heliconia unimaculata Hewitson, syntype <$ (Ecuador). 

Heliconius melpomene aglaope form niepelti Riffarth, syntype $ (Pastaza sup., 

Ecuad.). 

H. melpomene aglaope form iris Riffarth, holotype $ (Ob. Pastaza, Ecuad.). 

H. melpomene aglaope form isolda Niepelt, syntype ^(Jibaria). 

H. melpomene aglaope form cognata Riffarth, syntype $ (Pozuzo, Peru). 

H. melpomene aglaope form rubra Stichel, syntype <§ (Cuzco). 

H. aglaope Felder & Felder, syntype <£ (Rio Negro). 

H. melpomene aglaope form pavva Neustetter, holotype <§ (S. Columbien, Mocoa). 

H. melpomene aglaope form paula Neustetter, holotype <§ (S. Columbien, Mocoa). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 20 







204 




200 






'^■^205 





*M^206 





^207 




203 






208 



PLATE 21 
Upper- and underside 

Fig. 209. H eliconius melpomene aglaope form aurofasciata Neustetter, holotype $ (S. Columbien, 

Mocoa) . 
Fig. 210. H. amaryllis flagrans Stichel, syntype <-£ (Trinidad, Port of Spain). 
Fig. 2ii. H. melpomene eltvinghami Joicey & Kaye, holotype g (St. Laurent, Maroni River). 
Fig. 212. H. melpomene form luteipicta Neustetter, syntype £ (St. Laurent, Guyana). 
Fig. 213. H. melpomene form flavorubra Neustetter, syntype^ (St. Laurent, Guyana). 
Fig. 214. H. melpomene melpomene ab. primus Joicey & Kaye, holotype^ (Guyane Francse., 

St- Jean du Maroni). 
Fig. 215. H. melpomene melpomene ab. collis Joicey & Kaye, holotype g (Guyane Francse., 

Nouveau Chantier). 
Fig. 216. H. melpomene melpomene ab. compacta Joicey & Kaye, holotype^ (Guyane Francse., 

St-Jean du Maroni). 
Fig. 217. H. melpomene melpomene ab. melpina Joicey & Kaye, holotype £ (Guyane Francse., 

St-Laurent du Maroni). 
Fig. 218. H. melpomene melpomene ab. faivrei Joicey & Kaye, holotype 9 (Guyane Francse., 

St-Jean du Maroni). 



Bull. Br. Mus. nut. Hist. (Ent.) 32, 5 



PLATE 21 




PLATE 22 

Upper- and underside 

Fig. 219. Heliconins melpomene ab. atrosecta Riffarth, syntype <$ (Obidos). 

Fig. 220. H. melpomene funebvis form obscurata Riffarth, holotype <J (Berg en Dal, Sur.). 

Fig. 221. H. melpomene cybele ab. cybeleia Joicey & Kaye, holotype £ (Guyane Francse. 

St- Jean du Maroni). 
Fig. 222. H. melpomene cybele ab. negroida Joicey & Kaye, holotype <§ (Guyane Francse. 

St-Jean du Maroni). 
Fig. 223. H. melpomene cybele ab. fanstalia Joicey & Kaye, holotype ^ (Guyane Francse. 

St-Jean du Maroni). 
Fig. 224. H. melpomene form aurelia Neustetter, syntype <$ (Guyane Franc. Maroni). 
Fig. 225. H. melpomene cvbele ab. dianides Joicey & Kaye, holotype £ (Guyane Francse., 

St-Jean du Maroni). 
Fig. 226. H. melpomene form lavinia Neustetter, syntype $ (St. Laurent, Guyana). 
Fig. 227. H. melpomene cybele ab. elegantula Joicey & Kaye, holotype £ (Guyane Fr., St. Jean). 
Fig. 228. H. melpomene cybele ab. maris Joicey & Kaye, holotype § (Guyane Francse., 

St-Jean du Maroni). 



Bull. Br. Mus. mil. Hist. (Ent.) 32, 5 



PLATE 22 




226 




223 



PLATE 23 

Upper- and underside 

Fig. 229. Heliconius melpomene ab. diana Riffarth, holotype <£ (Cayenne?). 

Fig. 230. H. miitabilis Butler, syntype 9 (Amazons Trail). 

Fig. 231. H. melpomene thelxiope ab. stygianus Joicey & Kaye, holotype <-J (Guyane Francse. 

St- Jean du Maroni). 
Fig. 232. H. thelxiope var. aglaopeia Staudinger, syntype £ (Cayenne). 
Fig. 233. H. melpomene form penelopides Neustetter, holotype g (St. Laurent, Guyana). 
Fig. 234. H. melpomene thelxiope ab. negroidens Joicey & Kaye, syntype £ (Guyane Francse. 

St- Jean du Maroni). 
Fig. 235. H. melpomene form laurentina Neustetter, holotype g (St. Laurent, Guyana). 
Fig. 236. H. melpomene form athalia Neustetter, holotype g (St. Laurent, Guyana). 
Fig. 237. H. melpomene var. angusta Riffarth, paratype g (Cayenne). 
Fig. 238. H. melpomene thelxiope ab. majestica Joicey & Kaye, holotype $ (Guyane Francse. 

Nouveau Chantier). 



Bull. Br. Mus. uat. Hist. (Ent.) 32, 5 



PLATE 23 




229 





230 






236 



\ 




232 






237 




233 






238 



PLATE 24 
Upper- and underside 

Fig. 239. Heliconius melpomene thelxiope ab. lucindella Joicey & Kaye, holotype $ (Guyane 

Francse., St- Jean du Maroni). 
Fig. 240. H. melpomene thelxiope ab. punctavins Joicey & Kaye, holotype 9 (Guyane Francse., 

St- Jean du Maroni). 

H. liippolyte Bates, syntype <§ (Tapajos, Amazons). 

H. ritfolimbata Butler, syntype $ (Rio Tapayos). 

H. tyche Bates, syntype <J (Serpa, Amazons). 

H. melpomene madeira Riley, syntype $ (Lower Madeira). 

H. amaryllis var. euryades Riffarth, syntype^ (Vilcanota, Peru). 

H. penelope form aida Neustetter, syntype $ (Bolivia). 

H. penelope form excellens Neustetter, holotype £ (Bolivia). 

H. penelope form camea Neustetter, holotype £ (Bolivia). 



Fig. 


241. 


Fig. 


242. 


Fig. 


^43- 


Fig. 


244. 


Fig. 


^45- 


Fig. 


246. 


Fig. 


247- 


Fig. 


248. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 24 




239^^ I 





244 




248 



PLATE 25 

Upper- and underside 

Fig. 249. Heliconins penelope form noctis Neustetter, syntype $ (Bolivia). 

Fig. 250. H. penelope form amneris Neustetter, syntype <$ (Bolivia). 

Fig. 251. H. melpomene ab. margavita Riffarth, syntype £ (Bolivia). 

Fig. 252. H. amandus Grose-Smith & Kirby, syntype <£ (Bolivia). 

Fig. 253. H. phyllis form flammea Niepelt, holotype £ (Bolivia). 

Fig. 254. H. penelope form biedermanni Niepelt, syntype <J (Bolivia). 

Fig. 255. H. penelope form amandoides Neustetter, syntype $ (Bolivia). 

Fig. 256. H. nanna burchelli Poulton, syntype^ (Chapada, Matto Grosso). 

Fig. 257. H. melpomene burchelli form curvifascia Talbot, holotype g (Melguira, Matto Grosso). 

Fig. 258. H. melpomene burchelli form obscurifascia Talbot, holotype $ (Melguira, Matto Grosso). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 25 






255 




256 







257 




258 



Fig. 


259- 


Fig. 


260. 


Fig. 


261. 


Fig. 


262. 


Fig. 


263. 


Fig. 


264. 


Fig. 


265. 


Fig. 


266. 


Fig. 


267. 


Fig. 


268. 



PLATE 26 

Upper- and underside 

Heliconius nanna Stichel, syntype $ (Brasilia, Espirito Santo). 

H. galanthus Bates, syntype <§ (Forests of N. Vera Paz). 

Heliconia diotrephes Hewitson, syntype ^ (Nicaragua). 

Heliconius cydno galanthus form exornata Riffarth, syntype <J (Carillo, Costa Rica). 

H. chioneus Bates, holotype <§ (Lion Hill, Panama). 

Heliconia cydno Doubleday, syntype 9 (Bogota). 

Heliconius cydno cydnides form azteka Neustetter, holotype £ (Solano, W. Col.). 

H. cydno cydnides form confluens Neustetter, syntype <§ (Solano, Yapura). 

Heliconia aventina Oberthtir, holotype § (Nouve Grenade). 

H. temerinda Hewitson, syntype £ (Villagomes, N. Granada) . 






Bull. Br. Mus. not. Hist. (Ent.) 32, 5 



PLATE 26 




263 



PLATE 27 

Upper- and underside 

Fig. 269. Heliconia hermogenes Hewitson, syntype ^. 

Fig. 270. Heliconius zelinde Butler, syntype £ (W. coast of America). 

Fig. 271. H. cydno cydnides form flavidior Neustetter, holotype $ (Solano, Yapura). 

Fig. 272. H. cydno cydnides form albidior Neustetter, holotype £ (Rio Putumayo, Montegoa). 

Fig. 273. H. weymeri Staudinger, syntype^ (R. Dagua, Columb.). 

Fig. 274. H. weymeri ab. sulphureomaculata Fassl, syntype ^ (Rio Aguacatal, Colomb.). 

Fig. 275. H. weymeri submarginahis Fassl, syntype 9 (Rio Aguacatal, Colomb.). 

Fig. 276. H. weymeri ab. gustavi Staudinger, syntype $ (R. Dagua, Columb.). 

Fig. 277. H. wernickei Weymer, holotype^ (Bogota, Columbien). 

Fig. 278. H. emilius Weymer, holotype <$ (Muzo, Colombia). 



Bull. Br. Mits. nat. Hist. (Ent.) 32, 5 



PLATE 2 




269 











275 




278 



PLATE 28 



Fig. 


279. 


Fig. 


280. 


Fig. 


281. 


Fig. 


282. 


Fig. 


283. 


Fig. 


284. 


Fig. 


285. 


Fig. 


286. 


Fig. 


287 


Fig. 


288 



Upper- and underside 

Heliconius rubellius Grose-Smith & Kirby, syntype £ (Columb.). 

H. cydno broncus Stichel, syntype $ (Peru?). 

H. cydno ab. haenschi Riffarth, syntype <§ (Balzapamba, Ecuad.). 

Heliconia alithea Hewitson, syntype £ (Ecuador). 

Heliconius alithea neustetteri Riffarth, syntype <$ (Balzapamba, Ecuad.). 

H. cydno alithea form egregia Riffarth, syntype $ (Balzapamba, Ecuad.). 

H. melpomene timareta form insolita Riffarth, syntype <J (Santa Jnez, Ecuad. 

Heliconia timareta Hewitson, syntype $ (Ecuador). 

Heliconius pachinus Salvin, syntype $ (Chiriqui, Panama). 

H. hermathena form vereatta Stichel, holotype g (N. Brasilien, Faro). 



Bull. By. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 28 




PLATE 29 

Upper- and underside 

Fig. 289. Heliconius hermathena form hydarina Stichel, holotype $ (N. Brasilien, Faro) 

Fig. 290. Heliconia hermathena Hewitson, syntype $ (Amazons). 

Fig. 291. H. himera Hewitson, syntype <$ (Ecuador). 

Fig. 292. H. petiverana Doubleday, syntype $ (Mexico). 

Fig. 293. Heliconius hydara var. antigona Riffarth, holotype £ (Columbien). 

Fig. 294. Heliconia chestertonii Hewitson, syntype $ (N. Granada). 

Fig. 295. Heliconius molina Grose-Smith, syntype $ (Valdevia, Colombia). 

Fig. 296. H. hydara ab. nocturna Riffarth, holotype £ (Venezuela). 

Fig. 297. H. cyrbia ab. diformata Riffarth, syntype $ (Ecuador, Quito). 

Fig. 298. H. cyrbia cyrbia form bella Riffarth, syntype <$ (Slanos, Ecuador). 

Fig. 299. H. erato eratophylla Joicey & Kaye, holotype 9 (Tarapo, Peru). 

Fig. 300. H. erato estrella form agnata Stichel, holotype $ (Ucayali). 



Bull. Br. Mus. nut. Hist. (Ent.) 32, 5 



PLATE 29 








296 










294 





300 



PLATE 30 

Upper- and underside 

Fig. 301. Heliconius notabilis Salvin & Godman, syntype £ (Canelos, Ecuador). 

Fig. 302. H. erato simplex form rosacea Riffarth, holotype <§ (Pastaza sup., Ecuador). 

Fig. 303. H. microclea notabilis form radiata Oberthiir, holotype ^ (Guayaquil). 

Fig. 304. H. erato estrella from beata Riffarth, holotype 9 (Ob. Pastaza, Ecuad.). 

Fig. 305. H. erato estrella form rothschildi Niepelt, syntype £ (Ecuador, Canelos). 

Fig. 306. H. erato estrella form ochracea Riffarth, holotype^ (Ob. Pastaza, Ecuad.). 

Fig. 307. H. erato estrella form ilia Niepelt, holotype <J (Ob. Pastaza, Ecuad.). 

Fig. 308. H. etylus Salvin, syntype^ (Gualaquiza, Ecuador). 

Fig. 309. H. phyllis emma Riffarth, syntype g (Amazon super.). 

Fig. 310. H. erato estrella form palmata Stichel, holotype g (Ucayali). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 30 




301 






306 




302 





307 




303 






308 




304 




\ 





309 




305 






310 



PLATE 31 

Upper- and underside 

Fig. 311. Heliconius latlvitta Butler, syntype $ (Boa Vista, Rio Madeira). 

Fig. 312. H. estrella form glaucina Neustetter, holotype 9 (S. Columbien, Mocoa). 

Fig. 313. H. estrella form meliorina Neustetter, syntype $ (S. Columbien, Mocoa). 

Fig. 314. H. evato estvella form problemata Neustetter, syntype <£ (S. Columbien, Mocoa). 

Fig. 315. H. petiverana ab. tristis Riffarth, holotype $ (Chiriqui). 

Fig. 316. Heliconia hydara Hewitson, syntype $ (N. Granada). 

Fig. 317. Heliconius cyrbia var. juno Riffarth, holotype £. 

Fig. 318. H. phyllis var. viculata Riffarth, syntype $ (Surinam). 

Fig. 319. H . erato rubrizona Joicey & Kaye, syntype £ (Santarem). 

Fig. 320. H. phyllis ab. callista Riffarth, syntype $ (Amazon super.). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 31 





^€ 







317 




313 





314 






315^ V 





320 



PLATE 32 

Upper- and underside 

Fig. 321. Heliconius erato cybelinus form helena Riffarth, syntype g (Obidos). 

Fig. 322. H. phyllis ab. amalfreda Riffarth, syntype $ (Surinam). 

Fig. 323. H . phyllis ab. dryope Riffarth, syntype $ (Surinam). 

Fig. 324. H. coralii Butler, syntype £ (Serpa). 

Fig. 325. H. erato erato ab. cybelellus Joicey & Kaye, syntype $ (Serpa, Lower Amazon). 

Fig. 326. H. erato oberthueri Riffarth, syntype $ (Berg en Daal, Sur.). 

Fig. 327. H. erato ab. hemicycla Joicey & Kaye, syntype 9 (Guyane Francse., St. Jean du 

Maroni) . 
Fig. 328. H. erato erato form fuiginosa Riffarth, syntype 9 (Cayenne). 
Fig. 329. H. erato ab. constricta Joicey & Kaye, holotype £ (Guyane Francse., St. Jean du 

Maroni) . 
Fig. 330. H. vesta var. tellus Oberthiir, holotype g (Cayenne). 






Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 32 




321 



|jSr 





326 







327 








328 




324 






329 




325 






330 



PLATE 33 

Upper- and underside 

Fig. 331. Heliconius erato ab. telloides Joicey & Talbot, holotype $ (Guyane Francaise, 

Godebert, Maroni). 

Fig. 332. H . erato ab. protect Joicey & Kaye, holotype 9 (Guyane Francse., St. Jean du Maroni). 

Fig. 333. H. erato form roseoflava Neustetter, syntype $ (St. Laurent, Guyana). 

Fig. 334. H. estrella Bates, syntype <£ (Marajo I., Amazons). 

Fig. 335- H- phyllis andremona ab. androdaixa Seitz, holotype <§ (Bragance, Para). 

Fig. 336. H. erato estrella form simplex Riffarth, holotype £ (Cajon, Siid-Peru). 

F IG - 337- H- phyllis amphitrite Riffarth, syntype g (Hillapani, Peru). 

Fig. 338. H. erato amphitrite ab. unipuncta Joicey & Kaye, syntype <J (Tirapata, Perou). 

Fig. 339. H. venustus Salvin, syntype g (Apolobamba, Bolivia). 

Fig. 340. H. erato diva Stichel, holotype g (Bolivia). 

Fig. 341. Heliconia anactorie Doubleday, syntype g (Bolivia). 



Bull. Br. Mus. uat. His/. (Ent.) 32, 5 



PLATE 33 





337 




338 



333 



/ 




334 





339 




335 



1 i 





/ 




\\\\\ 



340 




336 




341 



PLATE 34 

Upper- and underside 

Fig. 342. Heliconius phyllis anactorie form henrici Krliger, holotype £ (Carmen, Nord Bolivia). 

Fig. 343. H. phyllis form leonora Kriiger, holotype $ (Rio Grande, Bolivia). 

Fig. 344. H. phyllis kruegevi Neustetter, holotype £ (Rio Grande, Bolivia). 

Fig. 345. H. phyllis anacreon form clelia Neustetter, holotype $ (Bolivia). 

Fig. 346. H. phyllis kruegeri from anacveonides Neustetter, holotype <$ (Rio Grande, Bolivia). 

Fig. 347. H. anacreon Grose-Smith & Kirby, syntype $ (Boliv.). 

Fig. 348. H. erato phyllis form difflnens Riffarth, syntype £ (Peru). 

Fig. 349. Papilio phyllis Fabricius, syntype <£. 

Fig. 350. Heliconius erato phyllis form sperata Riffarth, syntype £ (Cajon, Siid-Peru). 

Fig. 351. H. evato anacreon form anaitis Riffarth, holotype 9 (Prov. Sara, Siid Bolivia). 

Fig. 352. H. phyllidis Grose-Smith & Kirby, syntype <§ (Boliv.). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 34 




342 






347 





343 





344 




345 



^r^ "* T t^ 349 





351 




346 





352 



PLATE 35 

Upper- and underside 

F IG - 353- Heliconius erato ab. artifex Stichel, syntype £ (Brasilia, Espirito Santo). 

Fig. 354. H. octavia Bates, syntype £ (Duenas, Guatemala). 

Fig. 355. H. formosus Bates, syntype $ (Lion Hill, Panama). 

Fig. 356. Heliconia gynaesia Hewitson, holotype £. 

Fig. 357. H. longarena Hewitson, syntype £ (N. Granada). 

Fig. 358. Heliconius sotericus Salvin, lectotype $ (Guaymay, Ecuador). 

Fig. 359. H. telesiphe ab. nivea Kaye, lectotype $ (Perou). 

Fig. 360. Heliconia telesiphe Doubleday, lectotype $ (Bolivia). 

Fig. 361. Heliconius montanus Salvin, syntype £ (Orosi, Costa Rica). 

Fig. 362. H. clysonimus form flavopunctatus Fassl, syntype 9 (Rio Aguacatal, Columb. 



Bull Br. Mus. nat. Hist. (Ent.) 3J, 5 



PLATE 35 



■' 




/ 



353 





354 






358 




355 






359 




/ 



356 






! w 360 




357 






361 





362 



PLATE 36 
Upper- and underside 

Fig. 363. Heliconius clysonimns apicalis ab. semirubra Joicey & Kaye, syntype $ (Columbia, Rio 

Siato) . 

Fig. 364. H. clysonimus fische/i Fassl, holotype $ (Rio Aguacatal, Columb.). 

Fig. 365. Heliconia hygiana Hewitson, syntype 9 (Quito). 

Fig. 366. Heliconius charithonia punctata Hall, syntype $ (St. Kitts, B.W.I. ). 

Fig. 367. Eueides ricini insularis Stichel, holotype $ (Trinidad, Belmont). 

Fig. 368. Heliconia automatia Oberthur, holotype g (Guyane Francaise, St. Laurent du 

Maroni) . 

Fig. 369. Eueides egerifovmis Joicey & Kaye, syntype <£ (St. Jean du Maroni). 

Fig. 370. Heliconia eueidina Oberthur, syntype $ (Cayenne). 

Fig. 371. Heliconius demetev beebei Turner, holotype £ (British Guiana, Essequibo R.). 

Fig. 372. Eueides eratosignis Joicey & Talbot, syntype £. 

Fig. 373. Heliconius demetev turneri Brown & Benson, holotype £ (Ypiranga). 

Fig. 374. H. veraepacis Bates, syntype g (Forest of N. Vera Paz). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 36 




366 





369 





370 






v y 




372 





373 



368 





374 



PLATE 37 

Upper- and underside 

Fig. 375. Heliconius sava fulgidus Stichel, syntype £ (Carillo, Costa Rica). 

Fig. 376. Heliconia theudela Hewitson, syntype £ (Panama). 

Fig. 377. Heliconius magdalena Bates, syntype <§ (Bogota, Colombia). 

Fig. 378. H. sprucei Bates, syntype ^ (Chimborazo, Ecuador). 

Fig. 379. H. rhea ab. albinea Riffarth, holotype 9 (Surinam). 

Fig. 380. H. leucadia Bates, holotype^ (S. Paulo, U. Amazons). 

Fig. 381. H. antiochus ab. alba Riffarth, syntype £ (Amazonas sup.). 

Fig. 382. H. zobeide Butler, syntype <§ (Para). 

Fig. 383. H. sapho candidus Brown, holotype g (Ecuador). 

Fig. 384. H. congener aquilionaris Brown, holotype $ (Medina, Ost Colombia) 

Fig. 385. Heliconia eleuchia Hewitson, syntype ^ (N. Granada). 



Bull. Br. Mus. not. Hist. (Ent.) 32, 5 



PLATE 3; 




376^1 lw ^H f^fir 




377 









383 





384 




385 



PLATE 38 

Upper- and underside 

Fig. 386. Heliconius sapho primularis ab. deflava Joicey & Kaye, syntype ^ (Paramba, 

Ecuador) . 

Fig. 387. H. primularis Butler, syntype £ (Guayaquil). 

Fig. 388. Heliconia ceres Oberthiir, syntype $ (Nouvelle Granade, Cauca). 

Fig. 389. Heliconius cydno flaveola Joicey & Kaye, syntype £ (Venezuela, Mocotone). 

Fig. 390. H . aoede philipi Brown, holotype $ (Salampioni, Bolivia). 

Fig. 391. H. clysonymus tabaconas Brown, holotype £ (Charape, N. Peru). 

Fig. 392. H. melpomene timareta ab. virgata Stichel, syntype £ (Santa Jnez, Ecuador). 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 5 



PLATE 3! 






390 







392 



PLATE 39 

Upper- and underside 

Fig. 393. Papilio andremona Cramer, syntype £ (Surinam) 

Fig. 394. P. callycopis Cramer, syntype <$. 

Fig. 395. P. erythrea Cramer, syntype g. 

Fig. 396. P. amathusia Cramer, syntype £. 

Fig. 397. P. quirina Cramer, syntype £. 

Fig. 398. P. quirina Cramer, syntype §. 

Fig. 399. P. rhea Cramer, syntype J 1 (Surinam). 

Fig. 400. P. pasithoe Cramer, syntype $ (Surinam). 

Fig. 401. P. pasithoe Cramer, syntype £. 

Fig. 402. P. pasithoe Cramer, syntype £. 

Fig. 403. P. numata Cramer, syntype $ (Surinam). 

Fig. 404. P. silvana Stoll, syntype 9- 



Bull. Br. Mus. n-il. Hist. (Ent.) 32, 5 



PLATE 39 




ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177: 
18 plates, 270 text-figures. August 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 text -figures. September, 

1965- £3.25. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

£3.15. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae). Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text -figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text -figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 
68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. 
February, 1973. £6.60. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text-figures. December, 1973. £9.55. 

printed by Unwin Brothers Limited the gresham press old woking surrey England 



M ' I 



A REVISION OF THE GENUS 

PTYCHANDRA 

(LEPIDOPTERA : NYMPHALIDAE) 



H. J. BANKS, J. D. HOLLOWAY 

AND 

H. S. BARLOW 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 6 

LONDON : 1976 



A REVISION OF THE GENUS PTYCHANDRA 
(LEPIDOPTERA : NYMPHALIDAE) 



BY 

HENRY JONATHAN BANKS 

Division of Entomology, C.S.I.R.O., Canberra 

JEREMY DANIEL HOLLOWAY 

Tillinglea, Tillingbourne Park, Wotton, Dorking, Surrey RH5 6QL 

AND 

HENRY SACKVILLE BARLOW 

P.O. Box 295, Kuala Lumpur, Malaysia 



Pp. 215-252; 5 Plates; 22 Text-figures 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 6 

LONDON : 1976 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 6 of the Entomology 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent. 



ISSN 0514-6431 



Trustees of the British Museum (Natural History), 1976 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 10 March, 1976 Price £4.25 



A REVISION OF THE GENUS PTYCHANDRA 
(LEPIDOPTERA : NYMPHALIDAE) 

By H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 



CONTENTS 



Synopsis .... 

Introduction 
Acknowledgements 
Generic description and features 
Biology and habitat 
Biogeography and phylogeny 
Generic affinities 
Multivariate biometric study 
Hair pencil and brand structures 
Key to the species of Ptychandra 
Descriptions of the species . 
References .... 
Index ..... 



in Ptychandra 



Page 
217 
217 
218 
218 
220 
221 
224 
225 

234 
236 

237 
251 
252 



SYNOPSIS 

The genus Ptychandra (Lepidoptera : Nymphalidae, Satyrinae) is revised and a key to the six 
known species is given. P. negrosensis sp. n. and P. lorquinii leytensis subsp. n. are described. 
The distinctive male secondary sexual characters are described in detail. Biometric data are 
used to support the conclusions of qualitative taxonomic study, to place doubtful individuals 
and to discriminate a geographical race not separable by any single diagnostic character. The 
biogeographic and phylogenetic affinities of the genus and species are discussed. 



INTRODUCTION 

The genus Ptychandra Felder & Felder (Lepidoptera: Nymphalidae, Satyrinae) 
has long been in need of revision. The only works covering most of the genus 
are those by Fruhstorfer (1908; 1911) which are difficult to follow and often incorrect. 
The male brands and genitalia which are important diagnostic characters have not 
previously been illustrated. Capture by the authors (Barlow, Banks & Holloway, 
1971) while on the 1965 Cambridge Expedition to Mt Kinabalu, Sabah, Malaysia, 
of a series of the Bornean species, talboti, provided the stimulus for this work. 
Hitherto, with the exception of the unique holotype of talboti, the genus was only 
known from the Philippines. 

The material used in this revision was obtained from the British Museum (Natural 
History) (BMNH) ; the Hope Department of Oxford University (UM) ; the Sencken- 
berg Museum, Frankfurt-am-Main (SMN) ; the Humboldt Museum, Berlin (MNHU) ; 



218 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

the Carnegie Museum, Pittsburg (CM); the Smithsonian Institution, Washington 
(USNM); and the American Museum of Natural History, New York (AMNH). 
Additional invaluable material was obtained from the private collections of C. G. 
Treadaway and J. N. Jumalon whose Ptychandra were subsequently donated to 
the British Museum (Natural History). No Ptychandra were present in the Sabah 
Museum, Kota Kinabalu; the Raffles Museum, Singapore; or the Sarawak Museum, 
Kuching. 

Venation and cell nomenclature follows that in The Insects of Australia (C.S.I.R.O., 
1970), each cell being designated by the vein anterior to it. 

ACKNOWLEDGEMENTS 

We are deeply indebted to R. I. Vane- Wright and T. G. Howarth of the British 
Museum (Natural History) for much helpful advice and assistance in the early 
stages of the work and for criticism of the manuscript. H. Clench of the Carnegie 
Museum supplied information about the movements of the collector, Lorquin. 
J. N. Jumalon, University of San Carlos, Philippines and C. G. Treadaway gave us 
access to their private collections. H. Hayashi kindly sent us photographs of 
P. talboti specimens collected by a colleague on Mt Kinabalu in 1974. The collec- 
tion of the recent Bornean specimens was made possible through the generosity 
of a number of funds and companies which supported the 1965 Cambridge Expedi- 
tion to Mt Kinabalu. The statistical analyses of the biometric data were carried 
out by N. A. Campbell and A. Grassia of C.S.I.R.O., Division of Mathematics and 
Statistics. The photographs were taken by P. V. York (BMNH). 

GENERIC DESCRIPTION AND FEATURES 

PTYCHANDRA Felder & Felder 

Ptychandra Felder & Felder, 1861 : 304; Staudinger, 1887 : 222; Rober, 1889 : 203; Reuter, 
1897 : I22 » 553; Fruhstorfer, 1908 : 221; 191 1 : 329; Gaede, 1931 : 320; Miller, 1968 : 34, 
49; Hemming, 1967 : 388; Lewis, 1973 : 276. Type-species: Ptychandra lorquinii Felder & 
Felder, by monotypy. 

Description. <$. Fades. Upperside iridescent blue-purple with small white or 
bluish markings subapically and antemarginally. Underside light to deep brown ground colour 
marbled with darker brown, wavy transverse lines in discal area, with additional silver lines in 
some species. Full series of submarginal ocelli on hindwing underside in cells Rs to 1A -\- iA; 
that in cell Rs often enlarged and inwardly displaced. Forewing underside with two to four 
submarginal ocelli in cells Rs to M 3 . Venation (Text-fig. 1). Forewing cell approximately 
one-third length of costa. Vein M 3 colinear with lower discocellular vein. Median disco- 
cellular vein slightly angled at origin of M 2 , almost straight between origins of M 2 and M 3 , and 
meeting lower discocellular, being approximately at right angles. Origins of subcostal and 
cubital veins strongly swollen at base, anal vein somewhat less so. Hindwing cell closed, 
about half the length of the wing, with vein 5c + R 1 slightly inflated proximal to the humeral 
vein. Vein M 3 extended into a well formed tail with subsidiary lobes at terminations of 
CuA x and CuA 2 in some species. On hindwing, origins of CuA x and M 3 well separated. Secon- 
dary sexual characters. Forewing cubital branches displaced by a brand of mealy scales covered 
with a long black hair pencil placed between CuA x and M 3 (except schadenbergi) and a pouched 



REVISION OF GENUS PTYCHANDRA 



219 



hair pencil present basally between CuA 1 and CuA 2 (except talboti) running about one-third 
of length of wing in the interneural furrow. Hindwing hair pencil present in some species at 
origin of Rs overlying part of cell. Body and appendages. Eyes hairy. Palpi strongly haired 
on second segment with third segment approximately one-sixth length of second. Antennae 
about half length of costa with finely tapered club. Tibial spurs present. Genitalia (see 
Text-figs 2-14). Valvae sclerotized distally, usually with a three-lobed terminal section 
armed with flat sclerotized plates giving a heavily serrated tip. Exteriorly furnished with 
many narrow setae often as long as the breadth of the valva and interiorly possessing a few 
short stiff setae in the sclerotized zone. Tegumen and uncus smoothly tapered with the uncus 
often notched distally. Gnathos bifid, either smooth or with weakly sclerotized processes 
and plates on the dorsal surface distally. Posterior ends of gnathos curved upwards (except 
talboti). Aedeagus weakly sclerotized without cornuti. 

§. Fades. Upperside light brown with prominent white or off-white markings and broad, 
white, forewing subapical band. Underside with brown or whitish ground colour traversed 




hair pencil 
and brand 

hair pouch 



Sc+R, 



1A+2A 



M 3 y ^ CuA, CuA 2 

Fig. 1. Venation of Ptychandra lorquinii lorquinii. 



220 H. J. BANKS, J. D. H0LLOWAY & H. S. BARLOW 

by darker brown wavy lines in discal area. Submarginal ocelli similar to male but of increased 
size. Venation. Similar to male, with cubital branches displaced despite absence of brands 
or pouches. Body and appendages. Similar to male but antennae relatively shorter compared 
with forewing length. Genitalia. Outer margin of ovipositor lobes semicircular; posterior 
apophyses straight, about as long as radius of lobes. Bursa copulatrix equipped with two 
longitudinal rows of signa. 

BIOLOGY AND HABITAT 

The immature stages are undescribed. 

The Philippine species are woodland and forest butterflies found from sea-level 
to about 700 metres (with a unique record of lorquinii lorquinii from almost 2000 
metres). They are very much at home in the dappled light of glades and coppices 
in the forest, tending to avoid full, bright sunlight (Jumalon, in litt.). The males 
are often seen in numbers in small glades, settling on leaves in the shade and occa- 
sionally taking brief flights. Males have been observed to visit bird-droppings 
and overripe fruit and negrosensis, taken in hill rainforest with thick undergrowth 
of ferns and vines, was observed feeding from cut grass stems in the company of 
amathusiids and Neptis on an overcast day. In Luzon, lorquinii and leucogyne are 




Figs 2-5. Male genitalia of Ptychandra species. Left valve removed, aedeagus withdrawn 
and setation not shown. 2, lorquinii lorquinii; 3(a), schadenbergi ; 3(b), schadenbergi, 
aedeagus; 4, leucogyne, ex Luzon; 5(a), talboti, ex Mt Kinabalu, Borneo; 5(b), talboti, 
aedeagus. 



REVISION OF GENUS PTYCHANDRA 221 

segregated ecologically to some extent, the former being more common in advanced 
secondary woodland and the latter predominating in mature forest. 

In Borneo the Mt Kinabalu specimens of talboti were taken at about 2000 m 
close to primary forest along a road. The Trus Madi specimen was taken close to 
the summit in moss forest and the specimen of talboti from Mt Dulit was also cap- 
tured in montane forest. This species too is active in overcast weather. The 
flight habits of Ptychandra species appear similar. A female talboti was at first mistaken 
for a riodinid of a genus such as Zemeros, which is found in a similar habitat and 
has similar flight characteristics and colouration. The particular specimen sat 
on leaves at the forest edge with wings half opened in the sun, making short brisk 
flights between periods of rest. The larger members of the genus resemble some 
Lethe species in flight, wing pattern, colouration and habitat. There is no record 
of crepuscular flight for Ptychandra, but from this resemblance to Lethe it appears 
likely to occur. 

BIOGEOGRAPHY AND PHYLOGENY 

The genus Ptychandra is of great interest as it is one of very few butterfly genera 
that has undergone considerable radiation within the Philippine island group 









14 



Figs 6-14. Diagnostic details from male genitalia of Ptychandra. 6, gnathos of lorquinii 
plateni; 7, gnathos of lorquinii bazilana; 8, gnathos of lorquinii lorquinii; 9, tip of uncus, 
mindorana; 10, gnathos of mindorana; 11, tip of uncus, schadenbergi; 12, gnathos of 
schadenbergi; 13, gnathos of leucogyne; 14, gnathos of lorquinii leytensis. 



222 



H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

118° 1JP' 122° 124° 126° 




* Ptychandra lorqum 

T P. leucogyne 

+ P. schadenbergi 

m P. mindorana 

a P. talboti 

u P. negrosensis 



xlanac / 

\ x MI.Aroiot 1 

\ Mnnlolbon ^Polillo Is, 

BoloonT C-»* Quezon Cit> 
Mor,»eres^«'n'llfl ! A.p oele 
Pulonglugo--/* \xTCT„ s „ 




Fig. 15. Map of Philippines, Borneo and North Moluccas showing distribution 
records for Ptychandra species. 



REVISION OF GENUS PTYCHANDRA 223 

with little radiation elsewhere. Most of the Philippine butterfly fauna has been 
derived from mainland Asia and areas on the Sunda Shelf, especially Borneo (Hollo- 
way & Jardine, 1968; Holloway, 1973). Piychandra is a homogeneous genus with 
no close relatives and may therefore be considered monophyletic. Speculation 
on its radiation and speciation is limited by the paucity of material and patchiness 
of collecting. Some of the locality recording may be in error, such as the record 
of leucogyne from the North Moluccas (Halmaheira in the original description and 
Bat j an by Semper, 1889 : 60). The recent discovery of negrosensis suggests that 
other islands may yet yield further species. 

The Bornean species is morphologically very different from those in the Philippines 
(size, genitalia and secondary sexual characters) and could be regarded as an early 
offshoot within the genus. Of the Philippine species four are allopatric and the 
fifth, lorquinii, overlaps with leucogyne in Luzon and the central Philippines and 
with schadenbergi in the south (Text-fig. 15). Holloway (1973) suggested a general 
pattern of two phases of radiation with an intervening period of isolation and 
fragmentation, caused by changes in sea-level and therefore land area in the Pleisto- 
cene, to be responsible for the majority of present day butterfly distribution patterns 
in the East Indies. The distribution of Piychandra may be open to explanation 
along similar lines. The allopatric distribution of the Philippine species apart 
from lorquinii suggests the fragmentation of the range of a widespread common 
ancestor, followed by divergence of the fragments. P. mindorana, negrosensis 
and schadenbergi are probably still restricted to their original sites of isolation. 
P. leucogyne may have been isolated in the central Philippines and possibly the 
southern peninsula of Luzon, and lorquinii in Luzon. Given such a pattern of 
isolation, then the present distribution of lorquinii and extension north of leucogyne 
in Luzon would have resulted from a second spread phase. This is the simplest 
explanation of current distribution patterns within the genus. The areas of isolation 
correspond well with the distribution of land in the Philippines at periods of low 
sea-level as indicated by Darlington (1957), and the barriers indicated would no 
doubt have been intensified during periods of high sea-level. 

If more weight is placed on the secondary sexual differences of the males, especially 
those of the forewing hair pencil and associated scale patches, then an alternative 
hypothesis involving three spread phases could be considered. (1) spread of a 
generic ancestor over the Philippines and northern Borneo with isolation subsequently 
in three areas, leading to evolution of brand type (a) (see p. 234 for discussion of 
brand types) in Borneo, type (b) in the northern Philippines and type (c) in the 
south; (2) spread of the type (b) ancestor to Mindoro leading to the separation 
of mindorana and leucogyne, and spread of the type (c) ancestor north through the 
Philippines leading to the isolation and development of lorquinii on Luzon, negrosensis 
on Negros and schadenbergi (where the brand has been lost) on Mindanao; (3) spread 
of lorquinii south to Mindanao and leucogyne into the central Philippines. Further 
collecting in the central Philippines and observations of ecological differences 
between the species would help to clarify the picture. 

The subspecies of lorquinii, if one includes specimens from Samar and Bohol 
with leytensis (which they resemble most closely), correspond well with Semper's 



224 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

(1882) division of the Philippines into biogeographic provinces. The three males 
of lorquinii from Palawan are referable to /. plateni or I. bazilana rather than I. 
lorquinii and the species is not recorded from Mindoro. The Palawan form must 
have spread to Palawan via the Sulu Archipelago and north-east Borneo from 
Mindanao. The distribution of Eurema alitha (Felder), widespread in the Philip- 
pines and Sulawesi, extending down the Sulu Archipelago to north-east Borneo 
(Holloway, 1973), suggests that such spread would be possible. 

The ecological differences between lorquinii and leucogyne have already been 
mentioned and it is of interest that lorquinii, more characteristic of secondary 
growth, has proved the more dispersive, following the general pattern of greater 
mobility in secondary growth species indicated by Corbet (1941). 

The possibility of the occurrence of character displacement in lorquinii plateni 
and schadenbergi is discussed in the biometric section. 



GENERIC AFFINITIES 

The genus is of interest as it has proved difficult to place satisfactorily in the 
higher classification of the Satyrinae. Although usually regarded as an aberrant 
member of the Lethe group (Felder & Felder, 1861; Rober, 1888; Miller, 1968) the 
genus has characters linking it with other tribes. Miller (1968) suggested that the 
Melanitini gave rise to the Elymniinae and thus Lethini through steps resembling 
Ptychandra amongst others. Ptychandra has the origins of the hindwing veins 
CuA 1 and M 3 well separated. They are usually connate in the Elymniinae and 
particularly the Lethini. However, as noted by Fruhstorfer (1908; 191 1), many 
of the characters of Ptychandra are associated with the Mycalesis group rather than 
Lethe. In particular the prominently swollen bases of the forewing subcostal and 
cubital veins with the less prominent inflation of the anal vein base (Text-fig. 1) 
and the secondary male sexual characters are commonly found in the Mycalesini 
but not Lethini. There are structures very similar to the various brands, hair 
pouches and hair pencils of Ptychandra species found in representatives of Mycalesis 
and Orsotriaena. The location of these characters is unusual and is also Mycalesis- 
like rather than of the Lethini. The hindwing hair pencil at origin of Rs is also 
found in Mandarinia (Miller, 1968) and many Mycalesis (e.g. M. perseus) but not 
Lethe though several of that genus, e.g. Lethe minerva F., have a hindwing hair 
pencil arising just outside the cell with small modified scales beneath it. Reuter 
(1897) places Ptychandra closer to Neorina than Lethe but within his Lethidi on 
the grounds of the morphology of the palpi. 

Many features are consistent with a close relationship to the genus Lethe. The 
sexes are strikingly dimorphic with the female facies similar to species such as 
Lethe europa F. The wing shape with the slightly falcate forewing and prominent 
tailing at M 3 in the hindwing is lethine. Except as noted above, and where the 
secondary sexual wing characters cause displacement, the venation resembles 
Lethe, with the characteristic convexity and relative shortness of the forewing cell. 
This cell is extremely short in both male and female Ptychandra. As in Lethe 



REVISION OF GENUS PTYCHANDRA 225 

tibial spurs are present and the female foreleg is pentamerous with spines on the 
first four subsegments (Miller, 1968). 



MULTIVARIATE BIOMETRIC STUDY 

(written in conjunction with N. A. Campbell, 
C.S.I.R.O. Division of Mathematics and Statistics, Perth, Australia) 

The taxa in this study were previously defined on the basis of their differences 
in qualitative (discontinuous) characters, or on their locality of capture. The 
males of the lorquinii subspecies from Mindanao, Bazilan and Palawan are not 
readily separable from one another using qualitative (non-metric) characters. 
The separation of male lorquinii lorquinii from lencogyne is also difficult. 

On inspection there appeared to be a number of distinctive parametric (or quan- 
titative) characters related to the shape, structure and ornamentation of the wing 
which are possessed by the various taxa. A multivariate approach is adopted in 
this section to clarify the nature of the morphometric divergence in the wing 
characters between the taxa. 

Genetic divergence between taxa may be reflected in changes in the shape, 

Table i 

Biometric characters and ratios used for statistical analyses 

Character 

No. Character 

1 Forewing length. 

2 Forewing breadth. This is taken from the anal angle to the tangent to the 
costa, crossing the line from apex to origin of cubitus and radius at right angles. 

3 Hindwing length, taken from the inner angle of the cell to the maximum 
extent of the wing close to vein CuA l , not to the end of the tail at vein M 3 . 

4 Hindwing breadth, taken from termination of vein Rs to the maximum extent 
of the wing on the inner margin close to termination of \A + zA. 

5 The calculated ratio of the forewing breadth to length. This gives an index 
of relative narrowness of the wing. 

6 The calculated ratio of the hindwing breadth to length. This was designed 
to show the interspecific variation in hindwing shape. 

7 The perpendicular distance from the tip of the tail at vein M 3 , hindwing, 
to the line joining the maxium indentation in spaces A/ 2 and M 3 . 

8 Length of forewing lower discocellular vein. 

9 Length of forewing lower discocellular vein proximal to the origin of vein 
CuA j. 

10 Maximum length of hindwing cell. 

11 Distance of proximal displacement of centre of ocellus in forewing spaces 
M 3 (usually white-pupilled) from a line running through centres of ocelli in 
space M \ and M 2 . Distal displacements are recorded as negative values. 

12 Distal maximum width of outer yellow ring around ocellus in hindwing space 
Rs. 

13 Length of antenna. 

14 Maximum length of external hair pencil in space M 3 overlying brand. 



226 



H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 





Fig. i 6. Position of measurements taken for biometric study (see Table i for 
additional description) superimposed on venation of Ptychandra lorquinii lorquinii. 



REVISION OF GENUS PTYCHANDRA 



227 



structure and ornamentation of the wing. A multivariate study of the variation and 
covariation between characters is effective for elucidating subtle changes in shape 
and ornamentation. A small change in the ratio of two highly correlated characters 
between taxa provides a simple means of identifying the taxa; a multivariate 
approach (Blackith & Reyment, 1971) utilizing canonical variates potentially 
provides an even more effective means of separating and identifying taxa. Suites 
of correlated characters exhibiting subtle changes in their relative proportions 
between taxa can be combined to provide a final mathematical expression for 
identification. 

In order to study the divergence between the taxa, and particularly the problems 
discussed above, a series of biometric characters were measured as detailed in 
Table 1 and shown, where appropriate, in Text-fig. 16. Characters 1-4 were taken 
with a travelling microscope and characters 7-14 were measured with a microscope 
fitted with a calibrated eyepiece. All characters except 13 and 14 refer to wing 
shape, structure and ornamentation. 

The taxa considered in this study are given in Table 2, detailing numbers of 
individuals within the taxa for which a complete data series was available. The 
fragility of the specimens, most of which were collected more than 60 years ago, 
resulting in absence of tails or antennae, restricted complete data on some specimens, 
while scarcity (rarity) of collected individuals for other taxa restricted their 
numbers. The extensive series of leucogyne and lorquinii leytensis were obtained 
after completion of the biometric observations. 

The multivariate approach adopted to study the divergence between the taxa 

Table 2 

Symbols used and number of individuals measured for taxa studied biometrically. 





Symbol; 
text- figs 


for 
17-22 


No. of complete 
specimens meosured 




e? 


9 


tf 9 


P. leucogyne 


V 


T 


4 4 


P. lorquinii bozilona 


1 


© 


5 3 


P. lorqutnit leytensis 


\ 


® 


1 2 


P. lorquinii lorquinii 


* 


® 


10 6 


P. lorquinii plateni 


- 


e 


17 10 


P. lorquinii ex Palawan 


/ 


n.o. 


3 


P. mindorana 


D 


■ 


4 4 


P. negrosensis 


O 


♦ 


1 1 


P. schadenbergi 




ffl 


17 b 6 


P. talboti 


A 


A 


4 1 



a. taxa used for estimation of covanance matrices 

b. males without external hair pencil 
n.o. not applicable 



228 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 



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ow ring width 


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REVISION OF GENUS PTYCHANDRA 



229 



primarily utilizes canonical variates (cf. Blackith & Reyment, 1971). For many 
of the taxa there are too few specimens to define adequately the character means 
and covariance (or correlation) structure. Therefore it is not possible to examine 
formally if, as is assumed in a canonical variates approach, the character variances 
and correlations between characters are similar from taxon to taxon. Hence the 
approach adopted in this study is to examine the canonical variates for their taxo- 
nomic significance, as well as their apparent statistical significance. Canonical 
variates that do not provide useful taxonomic separation are not considered. 

A multivariate approach will only be of taxonomic value if the robustness of 
the results can be established. To this end, the covariance structure was determined 
from all the taxa, and from taxa with the largest numbers of specimens, namely 
lorquinii plateni, I. lorquinii and schadenbergi (males and females). Canonical 
analyses were determined with all taxa included, but with the covariance structure 
estimated by the two approaches detailed above. Alternatively, only the three 
taxa above were analysed; the remaining taxa were located by their canonical 
variate scores. Similarities in the relative affinities of the taxa, and the degree 
of separation between taxa, are taken to imply robustness of the results obtained. 
The results quoted here were produced by considering all taxa with the covariance 
structure estimated from the three longer series, since the three approaches gave 
comparable results with similar major groupings. 

Characters which do not contribute significantly to the discrimination between 




10-0 150 

Canonical variate I (m*10~ 4 ) 

Fig. 17. Biometric study. First and second canonical variate scores for all taxa. 



240 



230 



H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 



the taxa can be eliminated; these will generally be characters with small standardized 
canonical variate coefficients (i.e. the canonical variate coefficients multiplied by 
their respective pooled standard deviations). The effect of eliminating characters 
can be studied by observing the effect on the canonical root; little change means 
little loss of discrimination. 

An analysis of all taxa showed that only eight of the eleven characters contributed 
to the discrimination; further elimination of characters reduced discrimination 
between the taxa. The canonical variates and canonical roots (from the analysis 
with the covariance matrix estimated from the three taxa) for eight characters are 
given in Table 3. The canonical roots for the analysis with all characters included 
are also given for comparison. 

Individual scores for each taxon for the first two canonical variates are shown 
in Text-fig. 17. The first canonical variate reflects pronounced sexual dimorphism 
within the genus; this largely results from a contrast of character 9, the length of 
vein CuA v relative to character 8. The length of CuA 1 is strongly affected by the 
position of the brand in the male. 

The second canonical variate shows almost complete separation of the lorquinii 
subspecies from the rest of the genus. This separation results from a contrast 
between character 8, a measure of the forewing cell length, relative to the length 



12- 




P. lorquinii subspp. 



60 



100 



150 



200 



240 



Canonical voriole I (mxlO ) 

Fig. 18. Biometric study. Second and third canonical variate scores for all taxa. 



REVISION OF GENUS PTYCHANDRA 



231 



of the hindwing cell and forewing. Ocellus displacement, character 11, is also 
important. P. leucogyne and lorquinii lorquinii males are separated on this variate. 
The third canonical variate (Text-fig. 18) discriminates between talboti and the 
other taxa. 

The fourth canonical variate, when considered with the second (Text-fig. 19) 
achieves a remarkable clustering of the males and females of the individual taxa in 
this strongly dimorphic genus, confirming the correct pairing of the sexes. (The 
females of leucogyne were incorrectly taken to be the females of schadenbergi by 
Semper (1886). These two species are insufficiently distinguished to give proof of 
this error.) 

In the above analysis, the subspecific differences between the putative Palawan 
representatives of lorquinii and those from the other islands are masked because of 
the larger differences between lorquinii and the other species. A canonical analysis 
of the males of lorquinii from Luzon, Palawan, Mindanao and Bazilan (with the 



e 
H 

o 

o 




e \ 

"a e \ 



southern 
P. lorquinii subspp. 



<D/ 




P. negrosensis 



P. lorquinii lorquinii 



20-0 



240 



Canonicol variate H (mxlO ' 



Fig. 19. Biometric study. Second and fourth canonical variate scores for all taxa. 



232 



H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 



covariance structure estimated, as before, from the larger samples) was carried 
out. The results were again robust with respect to the approaches detailed above. 
The canonical variates and canonical roots are given in Table 4, while the canonical 
variate scores for all specimens for the first and second canonical variates are given 
in Text -fig. 22. The first canonical variate separates the Luzon representative 
(/. lorquinii) from the southern races, and to a lesser degree /. bazilana from I. plateni. 

Table 4 
Canonical variate scores for P. lorquinii subspecies (males only) using 9 characters. 



Character* 3 ) 










No. 


Character 


CV I 


CVII 


1 


Forewing length 


•050< b ) 


( 


77 o)W 


-•069 (-1-063) 


2 


Forewing breadth 


— -007 


(— 


083) 


•083 (-986) 


3 


Hindwing length 


•012 


( 


177) 


■070 (1-041) 


4 


Hindwing breadth 


— -oio 


(— 


148) 


— -077 (— 1-082) 


8 


Forewing cell length 


•016 


( 


187) 


•022 (-247) 


9 


Cell length to CuA j 


-•"3 


( — 


677) 


•127 (-762) 


10 


Hindwing cell length 


•058 


( 


538) 


-•047 (--437) 


11 


Ocellus displacement 


-•"3 


(— 


494) 


— •140 (— -610) 


12 


Yellow ring width 


-•194 


(-•305) 


-•210 (--33°) 




Canonical root 


1 


7i 


o-23 NS 




Canonical root from 










all characters 


1 


79 




0-25 



(a) See Table 1 

(b) Canonical variate score (for m x io~ 4 ) 

(c) Canonical variate score normalised for unit variance 



The second canonical variate (with the first) tends to separate the Palawan speci- 
mens from those of Bazilan and Mindanao. Analyses of the three taxa, with the 
covariance structure estimated by the approaches given above and from lorquinii 
subspecies only, all show some overlap of lorquinii bazilana and /. plateni, and 
separation of the individuals from Palawan and the other localities. While this 
separation may sometimes be inadequate for confident identification, it does 
indicate the likely taxonomic distinctness of the Palawan race from lorquinii plateni 
and I. bazilana. 

It is of interest to note that only the first canonical root is statistically significant. 
Clearly, there is marked overlap of the taxa along the second canonical variate; 
the means for /. lorquinii, I. bazilana and I. plateni are virtually coincident along this 
variate. And yet it is the second canonical variate which tends to separate the 
Palawan specimens from I. plateni and to a lesser degree I. bazilana. Hence both 
the taxonomic (or biological) and statistical significance of canonical variates should 
be examined in morphometric taxonomic studies. 

No information is available on ecological differences between lorquinii and schaden- 
bergi flying together in Mindanao but, given the sort of evolutionary history of the 
genus outlined above, /. plateni would appear to have undergone character displace- 
ment in Mindanao. Character displacement has been defined by Grant (1972) 



REVISION OF GENUS PTYCHANDRA 



233 



as 'the process by which a morphological character state of species changes under 
natural selection arising from the presence, in the same environment, of one or 
more species similar to it ecologically and/or reproductively'. In the case of Pty- 
chandra there is not the 'control' situation discussed by Brown & Wilson (1956) 
where both species, overlapping and interacting, exist also allopatrically so that 
comparisons can be made between the sympatric and allopatric states in each 
species. Only lorquinii occurs allopatrically in Luzon and the central Philippines 
(though interacting possibly with leucogyne). The suggestion that character 
displacement has occurred is based on the observation that in several characters 
where allopatric lorquinii lorquinii resembles schadenbergi, lorquinii plateni differs 
markedly from both. This is illustrated in several biometric parameters. P. 
lorquinii plateni is significantly larger than both other forms (P < o-ooi) indicated 
by characters 1-4, 10 and 13 (see Table 1). It differs significantly (P < o-ooi) 
in the displacement of the posterior forewing subapical ocellus character 11, plotted 
against character 1 in the scatter diagrams in Text-figs 20, 21. Whereas the white 
markings of the female forewing of /. lorquinii and schadenbergi are generally similar, 
those of /. plateni are modified into a band from the centre of the costa to the tornus 
as in female negrosensis. Biometrically (see Text-figs 16-19) lorquinii leytensis 
is placed with lorquinii plateni and lorquinii bazilana but in <$ and $ facies it is 




Fig. 20. Ocellus displacement against forewing length for male Ptychandra 
schadenbergi, P. lorquinii lorquinii and P. lorquinii plateni. 



234 



H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 



intermediate between these and lorquinii lorquinii. A clinal pattern cannot be 
ruled out. 

Positive identification of character displacement in plateni must await resolution 
of the second criterion identified by Grant (1972), that the observed differences 
between the sympatric and allopatric forms of lorquinii should be directly attribu- 
table to selection arising from the presence of schadenbergi. This awaits further 
study of the ecology of the species involved and of their variation. 

HAIR PENCIL AND BRAND STRUCTURES IN PTYCHANDRA 

The secondary sexual characters of male Ptychandra - brands, hair pencils and 
pouches - provide good diagnostic features at the species level and are of interest 
as one of the main differences between the genus and other Lethini. Consequently 
they have been studied in detail and their structures are given comparatively here 
rather than under each species. 

The most prominent features are found on the forewing just below the cell close 
to the origin of M 3 and are illustrated in Pis 1 and 2. Only schadenbergi does not 
possess a broad black hair pencil (PI. 2, fig. 4). For the remaining five species 
the pencil arises close to the origin of M 3 and usually covers a patch of broad modi- 
fied scales. Three forms of this structure are recognizable. 




Fig. 21. Ocellus displacement against forewing length for female Ptychandra 
schadenbergi, P. lorquinii lorquinii and P. lorquinii plateni. 



REVISION OF GENUS PTYCHANDRA 



235 



Type (a) (PI. i, fig. 1). The hair pencil arises from a poorly defined area at the 
base of cell M 3 which is characterized by brown, not purple, scales. The 
pencil overlays part of cell CuA x containing a roughly triangular area of slate- 
grey scales (talboti) . 

Type (b) (PL 1, fig. 2; PL 2, fig. 4). The hair pencil arises from a sharply defined 
pale elliptical area at the base of cell M 3 and extends over CuA v The vein 
is sunken there and the depression is filled with shiny black scales (leucogyne, 
mindorana) . 

Type (c) (PL 1, fig. 3; PL 2, fig. 6; Text-fig. 1). The hair pencil arises from the 
lower vein of the cell at the base of cell M 3 and extends into cell A/ 3 covering 
a well defined roughly circular area of pale mealy scales (lorquinii subspp. 
(except leytensis which lacks the mealy scales) , negrosensis) . 

In addition to these structures all species except talboti have a pouch in the inter- 
neural furrow of cell CuA 1 which runs from the base of the cell for about a third 
to half of its length. This contains a delicate light brown hair pencil of the same 
length as the pouch. The pencil arises external to the pouch at the origin of CuA 2 
and can be seen entering the pouch. It can be brought out by hooking a fine needle 
into the loop of hair between the pencil origin and where it enters the pouch. The 
structure is surrounded by an area of very small turquoise scales that are restricted 
to cell CuA x with forewing brand type (c) and surround the shiny black scales in 
brand type (b) . 

The hindwing brand structures (PL 3, figs 7-12) occur in the cell at the base of 



3C- 












I , Polowan subsp. ^\^ 




20- 








lorquinii 






/ ]\ 


10- 




X * 




X 




/ x\ / tS\_ 






X 






X 


/ ' 


1 / y 


plateni \ 


Q 






X 








_ | 








X 




X / 


~ ~ 


— 1 


1-0- 




1 






1 


— 1 — 


~~" 1 



14 16 

Canonical vanote I (m*10 ) 



Fig. 22. Biometric study. First and second canonical variate scores for 
Pty chandra lorquinii subspecies (males). 



236 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

Rs. In lorquinii no modification of the basic regular rows of purple scales is appa- 
rent. In mindorana there is an elongate area of larger, more irregular scales, while 
schadenbergi often has a concentration of hairlike androconia over a patch where the 
purple scales are lacking. In the other three species a prominent black hair pencil 
extends over a depression filled with black shiny scales at the origin of M v The 
general form is similar to that of forewing brand type (b). 



Key to the species of PTYCHANDRA 

1 Upperside predominantly deep blue. Males ....... 2 

- Upperside brown and white. Females ........ 7 

2 Black external hair pencil on forewing ........ 3 

- No such external hair pencil. 

Hair pouch between veins CuA x and CuA 2 present. Gnathos terminally hooked 

without dorsal teeth ....... schadenbergi (p. 244) 

3 Black hair pencil on upperside forewing arising at base of vein M 3 ... 4 

- Hair pencil arising in cell M 3 ......... 5 

^ Black hair pencil at anterior margin of hindwing cell. Two or less forewing sub- 
marginal ocelli. ......... negrosensis (p. 250) 

- No such hair pencil on hindwing. Three submarginal ocelli on underside forewing. 

lorquinii (p. 237) 

5 Forewing hair pencil arising from a well defined pale elliptical area and covering 

black specialized scales straddling CuA 1 . Forewing greater than 24 mm . 6 

- Forewing hair pencil arising from a loosely defined dark area and covering a triangu- 

lar patch of slate-grey mealy scales in cell CuA x . Forewing less than 21 mm. 
Four forewing submarginal ocelli. Gnathos smooth, without terminal hook 

talboti (p. 248) 

6 Black hair pencil in hindwing cell arising at base of vein Rs. Four forewing sub- 

marginal disjunct ocelli. Hindwing tail hooked. Gnathos smooth, tapered 
distally. Tip of uncus smoothly curved .... leucogyne (p. 246) 

- No such hair pencil. Three conjoined forewing submarginal ocelli. Tails of hind- 

wing straight. Gnathos toothed dorsally. Tip of uncus indented ventrally 

mindorana (p. 245) 

7 Discal area of upperside hindwing predominantly white or off-white ... 8 

- Discal area of upperside hindwing predominantly brown . . . . . 10 

8 Three conjoined submarginal ocelli on forewing underside. Tails at vein M 3 straight 

schadenbergi (p. 244) 

- Four disjunct submarginal ocelli on underside forewing. Tail at vein M 3 hooked . 9 

9 White area of forewing upperside not extending to inner margin. Forewing less than 

22 mm ........... talboti (p. 248) 

- White area of forewing upperside reaching inner margin. Forewing greater than 

24 mm. 

Tail hooked leucogyne (p. 246) 

10 Discal area of hindwing upperside with dull reddish area around costal ocellus. 

Forewing 35 mm negrosensis (p. 250) 

- Discal area brownish without reddening around costal ocellus . . . • J 1 

11 Hindwing tails straight, equal or subequal on veins M 2 , M 3 , and CuA x . 

Hindwing outer margin smoothly curved .... mindorana (p. 245) 

- Tail onJllj hooked and much longer than those on other veins. Tails on M 2 and 

CuA x much less than that on M 3 lorquinii (p. 237) 



REVISION OF GENUS PTYCHANDRA 237 

The brief key given by Fruhstorfer (191 1) for the separation of lorquinii and 
schadenbergi appears to be incorrect and misleading. P. schadenbergi is said to 
possess a hindwing hair pencil in the cell and a short stiff hair pencil below 3/ 3 
on the forewing. Both these features are found in leucogyne, only the latter in 
lorquinii and neither in schadenbergi. However, in the majority of specimens 
of schadenbergi there is a small narrow patch on the hindwing in the cell close to 
the origin of Ks where the hindwing hair pencil is found in other Ptychandra which 
lacks purple scales. Fruhstorfer states that lorquinii possesses a short hair pencil 
between '1st and 2nd median veins' (CuA 1 and CuA^. This may either refer to 
the long hair tuft enclosed in the deep pouch in this position or to the prominent 
short hair pencil arising in lorquinii close to the lower discocellular vein and shading 
a black androconial patch in cell M 3 . The extent of curvature of the lower disco- 
cellular vein used by Fruhstorfer (191 1) to separate schadenbergi and lorquinii 
is unreliable. 



DESCRIPTIONS OF THE SPECIES 

Ptychandra lorquinii Felder & Felder 
(Text-figs i, 2, 8; PI. i, fig. 3; PI. 3, fig. 7; PL 4, figs 13, 14) 

Ptychandra lorquinii Felder & Felder, 1861 : 304; Felder & Felder, 1867 : 498, pi. 68, figs 
1, 2, 3; Semper, 1886 : 58 (in part); Staudinger, 1887 : 22, pi. 78, 2 figs. 

Ptychandra lorqmni Fruhstorfer, 1899 : 80; 1908 : 222; 1911 : 330; Gaede, 1931 : 320; Lewis, 
J 973 : 2 ?6, pi. 172, fig. 1. [Unjustified emendation.] 

Description, q. Forewing length 22-9-29-2 mm. Fades. Upperside deep iridescent 
blue, diffusely margined with dark brown, sometimes extending to ocellar area. Forewing 
with subapical rhomboidal white fleck on costa and inwardly pointing submarginal white 
chevrons in cells Rs to lA with maximum convexity half way or more than half way distally 
on costa. The white markings may be reduced or diffuse in some subspecies. Forewing 
brand type (a) (PI. 1, fig. 1). Hindwing quadrate, tailed at M 3 , with cell M 2 excavated more 
deeply than cell M 3 , giving tail a hooked appearance. Tail obtusely terminated. Not 
tailed at vein M \. Hindwing brand in cell absent, hindwing ocelli sometimes darkly visible 
on upperside. Underside ground colour mid-brown with discal area traversed by three choco- 
late-brown wavy bands. Hindwing submarginal ocelli in cells Rs to \A disjunct, with that in 
cell Rs inwardly displaced and enlarged. Forewing with equal or subequal ocelli in cells 
M v M 2 and M 3 , that in cell M 3 displaced inwardly. Fringes white, interrupted with brown 
at vein ends. Antennae. Club tip rufous with base dark brown. Shaft with uniform grey- 
brown scales dorsally, rufous ventrally with pale flecks towards base. Genitalia. Valves with 
variable sclerotized processes at apex, long setae on outer surface. Aedeagus similar to that 
of schadenbergi (Text-fig. 3b). Gnathi with dorsal teeth and with tip hooked and broadened 
(e.g., Text-fig. 6). 

9. Forewing length 25-7-32-7 mm. Fades. Upperside forewing ground colour brown 
broken by a white subapical fleck, submarginal white chevrons, white spots in the position of 
the submarginal ocelli in cells M 2 and M 3 underside (which may be reduced or sometimes 
absent), and a broad white post discal band arising near centre of costa running diagonally to 
M 3 sometimes broken or conjoined with white subdiscal band extending from close to sub- 
marginal ocelli to below the cell but not reaching inner margin. The extension of the line of 
the forewing postdiscal band passes below white spot in cell M 2 if present. Underside forewing 



238 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

with three confluent ocelli decreasing in size posteriorly and with those in cells M 2 and M 3 
usually white. Wing shape similar to male but forewing costa evenly curved. Upperside 
hindwing discal area predominantly light brown but becoming off-white around ocelli in cells 
Rs to M 2 . Ocelli in cells CuA x and M 3 ringed with buff; those in cells Rs, M x , M 2 , \A + -zA 
darker, less distinct and confluent. Underside off-white ground colour marbled with brown 
with ocelli placed as male, edged inwardly and marginally with diffuse cream zone. Fringes 
off-white interrupted by light brown at vein ends. Antennae. As males but strongly flecked 
with pale scales basally. 

Distribution. Luzon, Marinduque, Leyte, Bohol, Samar, Panay, Mindanao 
Bazilan, Palawan. Recorded from the Polillo Is. (Fountaine, 1926) and Babuyan 
Is. (Semper, 1887) but these specimens were not found in the collections searched. 
Notably absent from material from Mindoro and Negros. 

Biology. See under subspecies of lorquinii. 

Discussion. Fruhstorfer (1899) and subsequent authors refer to lorquinii 
as 'lorquini\ an unjustified emendation. 

P. caerulans is considered here to be a synonym of lorquinii and is attributable 
to the nominate subspecies. The blue colour of the upperside, which is somewhat 
variable in lorquinii, is paler than usual in the unique male holotype of caerulans 
but otherwise it is not distinguishable. Genitalia are as lorquinii. Fruhstorfer 
(1908) himself expressed doubts as to the validity of the species. 

This species of Ptychandra exhibits a number of distinct island races, described 
below. 



Key to the subspecies of P. LORQUINII 

Upperside predominantly purple, males ........ 2 

Upperside brown and white or cream, females ....... 5 

White or brown mealy scales present beneath forewing hair pencil in cell M 3 . 3 

Mealy scales absent or scarce beneath forewing hair pencil. 

Forewing length 26-9-27-2 mm ..... lorquinii leytensis (p. 242) 

Hindwing underside ocelli white-pupilled, ringed outwardly with black, mid-brown, 

buff and mid-brown bands, the middle bands being separated by a thin dark 

brown line. Ocellus in forewing cell M 3 proximally displaced from the line joining 

those in cells M x and M 2 . 

Forewing length 22-9-27-8 mm .... lorquinii lorquinii (p. 239) 

Hindwing underside ocelli white-pupilled, ringed with black, light brown, buff and 

mid-brown without a dark dividing line between the middle bands. Ocellus in 

forewing cell M 3 either proximally or distally displaced ..... 4 

Forewing length 26-7-29-3 mm ..... lorquinii plat eni (p. 241) 

Forewing length 25-0-27-4 mm. 

Dorsal surface of gnathos with few but large teeth (Text-fig. 7) 

lorquinii bazilana (p. 242) 
Ground colour chocolate-brown. Subapical forewing white band constricted or 

broken in cell M 3 ....... lorquinii leytensis (p. 242) 

Ground colour sandy or reddish brown. Subapical forewing white band entire or 

broken in cell M 3 ........... 6 

Hindwing underside ocellus white pupilled and ringed outwardly with dark brown, 

buff, and mid-brown bands; the two buff bands being separated by a thin dark 



REVISION OF GENUS PTYCHANDRA 



239 



brown line. Forewing cell M 3 ocellus proximally displaced relative to those in 
cells M 1 and M 2 . 
Upperside subapical white band not continuous with discal white patch 

lorquinii lorquinii (p. 239) 

- Hindwing underside ocellus white pupilled and ringed with dark brown, light brown, 

buff and mid-brown bands, without a dark dividing line between the two middle 
bands. Forewing cell M 3 ocellus distally displaced relative to those in cells M x 

and M 2 7 

7 Broad unbroken postdiscal off-white band running from halfway along costa to 

discal white area, sometimes slightly constricted in cell M 3 lorquinii plateni (p. 241) 

- Subapical off-white postdiscal band strongly constricted or broken in cell M 3 

lorquinii bazilana (p. 242) 

The key above has been based where possible on qualitative rather than bio- 
metric characters. However, in the case of male lorquinii plateni and /. bazilana 
differentiation is difficult and may be assisted by measurement of several parameters 
(see biometric section). The range of measurements given is that actually observed 
and is limited by the sample size (see individual descriptions) . 



Ptychandra lorquinii lorquinii Felder & Felder 
(Text-figs i, 2, 8; PI. 1, fig. 3; PL 3, fig. 7; PI. 4, figs 13, 14) 



Ptychandra lorquinii lorquinii Felder & Felder, 1861 : 304. LECTOTYPE $, Philippines: 

Luzon (BMNH), here designated [examined]. 
Ptychandra caerulans Fruhstorfer, 1908 : 223; 191 1 : 330; Gaede, 1931 : 320. LECTOTYPE 

(J, Philippines: Luzon (BMNH), here designated [examined]. Syn. n. 
Ptychandra lorquinii f. obscurior Fruhstorfer, 1908 : 223; Gaede, 1931 : 320. Type-material 

(not definitely located), Philippines: Luzon. Syn. n. 
Ptychandra lorquinii obscurior Fruhstorfer; Fruhstorfer, 191 1 : 330. 

Description. ,$. Forewing length 22-9-27-8 mm, n = 15. Fades. Upperside deep 
iridescent blue with forewing prominently marked with a white apical fleck and submarginal 
chevrons. Underside usually traversed with metallic wavy bands. Hindwing ocelli with 
white pupils ringed outwardly with black, mid-brown, buff and mid-brown bands, the middle 
bands being separated by a thin dark brown line. Genitalia (Text-fig. 6). Gnathos with 
broadened, hooked distal ends and two or more prominent dorsal teeth. 

$. Forewing length 25-7-28-4 mm, n = 8. Fades. Upperside ground colour mid-brown 
with a subapical white band extending from vein M 3 to approximately halfway along costa, 
usually disjunct from subdiscal white band but occasionally just meeting it. Subdiscal white 
band usually running almost completely along the length of inner margin but separated from 
it by a thin brown area. White spot corresponding to cell M 2 underside ocellus present and 
often a similar spot corresponding to cell M 3 . Width of yellow ring of hindwing underside 
costal ocellus 0-4-1-0 mm, x = 0-76 mm, n = 16. Ocelli white-pupilled, ringed successively 
with dark brown, buff, buff and mid-brown bands; the two buff bands being separated by a 
thin dark brown line. 

Distribution. Luzon. Five males attributable to this subspecies in Carnegie 
Museum were labelled 'Mindanao'. As specimens from other genera from the same 



240 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

collection were labelled Mindanao, yet of the Luzon subspecies (H. Clench, personal 
communication), it seems reasonable to assume that these Ptychandra specimens 
were mislabelled. 

Biology. Said to fly from April to January (Semper, 1886) but material has 
been seen from almost all months. 

Discussion. The holotype of P. caerulans is attributable to this subspecies. 

P. lorquinii obscurior is synonymized here. It was described (Fruhstorfer, 1908) 
as having the white areas of the hindwing reduced, a normal feature of the typical 
subspecies. First described as a forma, it was later elevated to subspecific rank 
(Fruhstorfer, 191 1) without comment. An apparently normal female from Fruh- 
storfer's collection bears his 'type' label (see Material examined below) and corres- 
ponds with his brief description. He refers for differentiation to the illustration 
given by Staudinger (1892), but this is not substantially different in marking from 
the lectotype.. The type of this form or subspecies did not appear in the catalogue 
of the Fruhstorfer collection (Martin, 1922 : 66). In the description of the female 
forma obscurior (Fruhstorfer, 1908) the specimens, presumably males, are described 
with androconial patches as for the typical form and are not distinguished. 

In addition to variation in colour of the purple upperside lorquinii varies consider- 
ably in the brightness of the underside markings and the distinctness of the sub- 
apical fleck and antemarginal chevrons. The underside of some specimens is 
marked with metallic wavy lines, as commonly found in mindorana. One specimen 
has been found with two forewing ocelli and one with four, the extra spot being in 
cell CuA lt not cell R 5 as in leucogyne (q.v.). 

Material examined. 

Ptychandra lorquinii, lectotype <$, Luzon, genitalia preparation no. TGH 1966-689 
(BMNH); 1 $ paralectotype, Luzon (Lorquin) (BMNH); 3 g putative paralectotypes, 
Luzon (BMNH). Ptychandra caerulans, lectotype <$, Luzon, genitalia preparation 
no. TGH 1966-690 (BMNH). Ptychandra lorquinii f. obscurior, syntype $, Luzon 
(BMNH). 

Luzon: 16 km W. of Baguio, 2500 ft, 1 $, 14. xi. 1966 (C. G. Treadaway) (BMNH); 
Baguio, 2 (J, 2 $, (CM); Baguio City, 3 $, ix. 1957 (C. G. Treadaway) (BMNH); 
Bataan, 1 <$, 11-20.vi.1880 (BMNH); Cape Engano, 1 <$ {Whitehead) (BMNH); 
Klondyke, 800 ft, 19 <$, 6 $, 19-27. xii.1911, 1-26.iv.1912, 23.V.1912 {A. E. Wileman) 
(BMNH); Lepanto, 1 $ {Whitehead) (BMNH); Los Banos, 1 J, 10.ix.1920 (BMNH); 
Los Banos, 1 $, 10. ix. 1920 (AMNH); Los Banos, 1 <$, 14. v. 191 1 {A. E. Wileman) 
(BMNH); Los Banos, 1 $, 3 $, (BMNH); Marivelis, 1 $, 11.xi.81 (SMN); Montalban, 
Rizal, 1 <$, 2 $, 4. i, 5.11.1914 {A. E. Wileman) (BMNH); Montalban, 1 <$, 29.viii.1966 
(C. G. Treadaway) (BMNH); Montalban 100 m, 1 J, 26.vi.1971 (C. G. Treadaway) 
(BMNH); Mt Arajat, 1 <J, 1 $, viii. 1903 {Browne) (BMNH); Mt Makiling, 1 <$, 
4.viii.i956 (C. G. Treadaway) (BMNH); Mt Makiling, 1500 ft, 1 <$, 3 $, viii, 4.ix.i966 
(C. G. Treadaway) (BMNH); Palali, Benguet, 2000ft, 6 <$, 5 $>, 21-25. xiii.1912, 
27.vi.1913 {A. E. Wileman) (BMNH); Pulang Lupa, Rosales, 1 $, 12.xi.81 (SMN); 
Quezon Nat. Park, 1100ft, 3 $, 2 $, 27-29. iv.1969 (C. G. Treadaway) (BMNH); 



REVISION OF GENUS PTYCHANDRA 241 

Quezon Nat. Park, Atimonan Road, i <$, 18.V.1968 (C. G. Treadaway) (BMNH) ; 
Sierra Madre, 1 <J (C G. Treadaway) (BMNH); Tarlac, 4 <J {Browne) (BMNH); 
Tagaytay Ridge, 1800 ft, 9 <$, 8 $, 3.ix.i966 (C. G. Treadaway) (BMNH); 'N. Luzon' 
5000-6000 ft, 1 S [Whitehead) (BMNH); Luzon, 1 <J, 2 $ (BMNH). Marinduque: 
3 (J, 1 $, vi. 1972, x. 1973 (C. G. Treadaway) (BMNH). Mindanao: Mindanao, 
5^ {Haslam) (CM). 

Ptychandra lorquinii plateni Semper stat. n. 
(Text-fig. 6; PI. 4, fig. 15) 

Ptychandra lorquinii Felder & Felder; Semper, 1886-95 [ m part]. 

Ptychandra plateni Semper, 1892 : 328. LECTOTYPE <$, Philippines: North Mindanao 
(SMN), here designated [examined]. 

Ptychandra lorquinii mindanaensis Fruhstorfer, 1899 : 80; 1908 : 223; 1911 : 330. LECTO- 
TYPE $, Philippines: Mindanao (BMNH), here designated [examined]. Syn. n. 

Ptychandra lorquinii var. mindanaensis Fruhstorfer; Gaede, 1931 : 320. 

Description. $. Forewing length 25-7-29-2 mm, n = 22. Fades. Upperside dull iri- 
descent blue narrowly margined with dark brown. White apical fleck and marginal white 
chevrons faint or absent. Turquoise scales scattered between discal cell and costa. Under- 
side marbled with dull brown and light brown without metallic wavy bands. Marginal ocelli 
with bluish white pupil ringed with black, light brown, buff and mid-brown, without a dark 
thin dividing line between the middle rings. Forewing submarginal ocellus in cell M 3 either 
proximally or distally displaced (0-9-1-0 mm, x = 0-02 mm, n = 25). Genitalia. Similar to 
those of lorquinii lorquinii (Text-fig. 2) but gnathos tending to be narrower at posterior end, 
less strongly reflexed and having more but smaller dorsal teeth which extend further anteriorly 
(Text-fig. 6). Valves similar to those of lorquinii lorquinii (Text-fig. 2) but lacking central 
terminal process. 

9. Forewing length 28-5-32-7111111, n = 11. Fades. Broad unbroken median white band 
on forewing upperside running from about halfway along costa to tornus and with the subdiscal 
area uniformly brown. Median band sometimes slightly constricted in cell M t . Submarginal 
white spot in cell M 2 present, in cell M 3 absent on upperside forewing. Ground colour sandy 
brown. Forewing submarginal ocellus in cell M 3 displaced distally (0-1-1-3 mm, x = 0-74 mm, 
n = 18). Maximum width of mid-brown band on costal ocellus, 0-5-1-0 mm, x = o-68, 
n = 18. Ocelli with white pupil ringed with dark brown, light brown buff and mid-brown 
bands. Genitalia. Bursa copulatrix with signa consisting of two parallel bands of spines dorsally, 
each consisting of about 100 rows, length 2-5 mm, of straight short teeth placed 3 abreast. 
Width of each rank of 3 about 0-2 mm. 

Distribution. Mindanao. One specimen found labelled 'Manille'. 

Biology. Said to fly from May to October and again in December and January 
(Semper, 1892). 

Discussion. Fruhstorfer (1899) considered plateni Semper to be a nomen nudum 
but Semper (1892) in his brief description refers back to his description of P. lorquinii 
in which the races of Luzon and Mindanao are adequately distinguished. The 
name plateni is therefore valid. 

One male example of this subspecies was found which, though normal in pattern 
and structure, was abnormally large, having a forewing length of 31-0 mm. 



242 h. j. banks, j. d. holloway & h. s. barlow 

Material examined. 

Pty chandra lorquinii plateni, lectotype <$, Mindanao: North, coll. C. Semper 
'Typus SMFL 20' (SMN) ; paralectotypes, 1 <$, 1 9, N. Mindanao (SMN) ; 1 J, 12.L1882, 
Sibulan (SMN); 1 <J, Bullonis, S., (SMN); 3 <$, Mindanao (BMNH); putative 
paralectotypes, 5 <J, 3 9, Mindanao {Platen) (MNHU); 1 $, Davao {Platen) (MNHU). 
Ptychandra lorquinii mindanaensis , lectotype 9> Mindanao (BMNH); paralectotype 
<J, Mindanao (BMNH). 

Mindanao: Agusan, Davao, 1 3, v. 1959 (/. N. Jumalon) (BMNH); Calian, 
Davao, 1 $, 31.V.1930 (C. F. Clegg) (CM); Davao, 2 ',i? (P/aten) (BMNH); Davao, 
2 ^, 1 9> iii. 1890 (BMNH); Jaliobong, Agusan, 1 $, 15.V.1968 (C. G. Treadaway) 
(BMNH); Kolambugan, Lanac, sea level, 1 $, 20.V.1914 (.4. E. Wileman) (BMNH); 
Margosatubi, 1 £, (BMNH); Samboangan, 1 9, ix. 1891 (BMNH); Talaud, Davao, 

1 $ {0. Jumalon) (BMNH) ; Talaud, Davao, 1 <$, 8.ii.i96i (C. G. Treadaway) (BMNH); 
Mindanao, 1 $ (/. /. Mounsey) (BMNH) ; Mindanao, 16 $, 5 9, 1903-1904 (/. Water- 
stradt) (BMNH); N. Mindanao, 1 $ (BMNH); 1 <J {Brabant) (BMNH); Mindanao, 

2 S, 3 $ (BMNH). " Luzon: 'Manille', 1 $, (BMNH). 

Ptychandra lorquinii bazilana Fruhstorfer 
(Text-fig. 7; PL 4, fig. 16) 

Ptychandra lorquinii bazilana Fruhstorfer, 1899 : 79; 1908 : 223; 1911 : 330, pi. 93, row g, 

second fig. LECTOTYPE $, Philippines: Bazilan (BMNH), here designated [examined]. 
[Ptychandra caerulans Fruhstorfer; Fruhstorfer, 191 1 : pi. 93, row g, first fig. Mislabelled.] 
Ptychandra lorquinii var. bazilana Fruhstorfer; Gaede, 1931 : 320. 

Description. <$. Forewing length 25-0-27-4 mm, n = 6. Fades. As in lorquinii plateni. 
Genitalia. Similar to those of lorquinii plateni but with apex of gnathos less broadened, less 
reflexed and with a few but more prominent dorsal teeth (Text-fig. 7). 

9- Forewing length 27-8-30-1 mm, n = 3. Fades. Forewing postdiscal white band con- 
stricted or broken in cell M 3 with slight interruptions produced by dusting of black scales 
along M 3 and CuA v Otherwise similar to lorquinii plateni. Forewing submarginal ocellus in 
cell M 3 weakly displaced outwardly (0-1-0-7 mm, x = 0-30 mm, n = 4). Outer mid-brown 
band on hindwing costal ocellus 0-3-0-6 mm, x = 0-48 mm, n = 4. 

Distribution. Bazilan. 

Biology. Not recorded. 

Material examined. 

Ptychandra lorquinii bazilana, lectotype 9, Bazilan, ii-iii. 98 {Doherty) (BMNH). 
4 3, 2 9 paralectotypes, Bazilan, 11— iii. 98 {Doherty) (BMNH); 1 g paralectotype, 
Bazilan ii-iii. 98 {Doherty) (CM). 

Bazilan: 16 £ (BMNH); 1 <J (C. /. Grist) (BMNH). 

Ptychandra lorquinii leytensis subsp. n. 

(Text-fig. 14; PI, 4, fig. 17) 
Description, g. Forewing length 26-9-27-2 mm. Fades. Slightly larger than nomino- 



REVISION OF GENUS PTYCHANDRA 243 

typical form and more boldly marked. Upperside markings as in lorquinii lorquinii but with 
distinct, pale blue-white marginal line on upperside hindwing. Forewing brand type (a) (PI. 1, 
fig. 1) but with mealy scales beneath the hair pencil much reduced or absent. Underside with 
dull silver streaking but otherwise markings as in plateni. Genitalia. Similar to those of 
lorquinii plateni, except gnathos less strongly hooked and dorsal teeth on it weakly developed 
(see Text-fig. 14). 

9- Forewing length 28-7-30-4 mm. Fades. White markings resemble those of lorquinii 
bazilana but with pinkish tinge in white forewing postdiscal band and no dusting of black 
scales along CuA x and M 3 . Ground colour of upperside dark chocolate brown. Forewing 
submarginal ocellus in cell M 3 displaced outwardly. Hindwing costal ocellus, outer mid-brown 
ring broad (o-8-i-i mm, x = i-o mm, n = 6). 

Distribution. Leyte. 

Biology. Found quite commonly in forest clearings up to 700 m (/. N. Jumalon, 
personal communication). 

Discussion. This subspecies is distinguished in the female from other lorquinii 
by the different shade of brown markings. The white markings can be regarded 
as transitional between the nominotypical species of Luzon and lorquinii plateni 
from Mindanao. The male is similarly intermediate, having the larger size of the 
Mindanao form with the usually more bold markings of the Luzon race. The 
forewing brand is distinctive with its reduced mealy scaling below the hair pencil. 

Material examined. 

Holotype <$, Leyte: Catamon, St Bernard, 800 ft, 6.vi.i967 (0. Jumalon) (BMNH). 

Paratypes. Leyte: Catamon, St Bernard, 3 $, 6, vi., 6.xi.ig67 (0. Jumalon) 
(BMNH); Lake Danao, Ormoc, 1 $, 29.ix.1963 (J. N. Jumalon) (BMNH); Lower 
Tangunan, Ormoc, 1 $, 21. xi. 1963 (/. N. Jumalon) (BMNH); Tungunan Hot Springs, 
Cananga, 2 <$, 20. xi. 1966 (J. N. Jumalon) (BMNH). 

Non-paratypic material. Leyte: Catamon, St Bernard, 1^,1$, vi. 1967 (/. N. 
Jumalon) (BMNH); Lake Danao, Ormoc, 1 $ {J. N. Jumalon) (BMNH); Ormoc, 
2 <$, 10.viii.1965, 10.xi.1965 (/. N . Jumalon) (BMNH); Ormoc, 1^,1$, (C. G. Tread- 
away) (BMNH); Ormoc, 1 j, 3.X.1965 (J. N. Jumalon) (BMNH); Tungunan Hot 
Springs, 2 <$, 1 9, 17.xi.1968, xi. 1966, (J. N. Jumalon) (BMNH); West Leyte, 1 <$, 
xi. 1966 {J. N. Jumalon) (BMNH). 



Ptychandra lorquinii subspp. incertae sedis 

Occasional specimens of lorquinii have been seen from Palawan, Bohol, Panay 
and Samar. The Palawan specimens, all of which are males, are not distinguishable 
from lorquinii plateni except biometrically (see biometric section). The lower 
discocellular vein on forewing of lorquinii bazilana is significantly shorter than 
from the Palawan specimens and they are distinct from the Luzon specimens. 
The male specimens from Panay and Samar closely resemble lorquinii leytensis 
in their prominent white upperside markings, relatively broad hindwings and 



244 H - J- BANKS, J. D. HOLLOWAY & H. S. BARLOW 

strongly coloured undersides with dull silver streaking. The female specimens 
from Bohol (see PI. 4, fig. 18) and Samar closely resemble lorquinii plateni and 
lorquinii leytensis in patterning, particularly in having a similar subapical white 
band and similarly ringed ocelli. When a longer series of these insects has been 
taken from these islands it is likely that subspecific differences will become apparent. 

Material examined. 

Bohol: i $ (SMN). Palawan: i $ {Platen) (BMNH); 1 J (BMNH). Panay: 
Yiollo, 1 S, iv. 28 (USNM). Samar: Antiago Valley, 1 $ (/. /. Mounsey) (BMNH); 
1 $, vii-viii. 1896 (/. Whitehead) (BMNH). 

Ptychandra schadenbergi Semper 
(Text-figs 3a, b, 11, 12; PL 2, fig. 4; PL 3, fig. 8; PL 4, figs 19, 20) 

Ptychandra schadenbergi Semper, 1886 159, pi. 11, fig 6; Reuter, 1897 : 122; Fruhstorfer, 
1899 : 80; 1908 : 222; 191 1 : 329, pi. 93, row g, third fig.; Gaede, 1931 : 320; Hobby, 1940 : 
220; Lewis, 1973 : 276, pi. 172, fig. 2. LECTOTYPE $, Philippines: North Mindanao 
(SMN), here designated [examined]. 

Ptychandra schadenbergi f. hebetatrix Fruhstorfer, 1908 : 222; 191 1 : 329; Gaede, 1931 : 320. 
LECTOTYPE $, Philippines: Mindanao (BMNH), here designated [examined]. 

Description. <§. Forewing length 23-2-26-7 mm, n = 23. Fades. Upperside deep iri- 
descent blue with prominent white submarginal chevrons and white apical fleck on forewing. 
Forewing ocelli in cells M lt M 2 and M 3 and those in M 2 and M 3 confluent. Fringes brown. 
Pouched hair pencil present in cell CuA 1 forewing below cell containing long dark brown hairs 
and narrowly bordered with turquoise scales. Forewing subquadrate with maximum curvature 
distal of centre of costa. Hind wing outer margin evenly curved with well defined scalloping 
between veins. Hindwing tail on M 3 short, straight, weakly hooked and terminated acutely. 
Underside ground colour brown with darker marbling. Hindwing with usual ocelli with white 
pupil ringed by black, dark brown, buff and dark brown. Hindwing hair pencil in cell absent 
but small patch of modified dark scales present close to origin of Rs in most specimens. Anten- 
nae. Rufous, darker dorsally, especially on club. Thin cream line ventrally extending from base 
to two-thirds length of antennae. Genitalia (Text-fig. 3a, b). Tip of uncus slightly concave 
(Text-fig. 11). Gnathi (Text-fig. 12) without dorsal teeth, becoming distinctly narrower just 
before terminal hook. Valves tending to have more than one main terminal process. 

9- Forewing length 25-7-28-8 mm, n = 9. Fades. Upperside mid-brown with white 
markings. Wing shape similar to male. Forewing with rhomboidal white subapical fleck on 
costa, white submarginal ocellar spot in cell M 2 , white postdiscal band running from costa 
towards spot in cell M 2 . White discal patch bounded on inner margin by brown, never meeting 
this edge. Hindwing discal area white, bounded outwardly by brown, inward of the ocellar 
area. Ocelli sometimes faintly visible on upperside hindwing, particularly in cell CuA 2 . White 
markings on underside as upperside but traversed by wavy brown thin bands in discal area. 
Forewing with three conjoined ocelli in cells M lt M 2 and M 3 . Ocelli white pupilled, distinc- 
tively ringed in succession with black, dark brown, buff and dark brown. Antennae. Patterned 
as male. 

Distribution. Mindanao. One specimen each examined labelled Palawan 
and Luzon, possibly with locality in error. 

Biology. Not recorded. 



REVISION OF GENUS PTYCHANDRA 245 

Discussion. A female specimen of leucogyne was illustrated and described by 
Semper (1886) as the female of schadenbergi, although Semper had female specimens 
of schadenbergi in his collection dated before his published description (cf. 1 $, Sibulan, 
i.xii.81) and possibly even from the same collector (the illustrated specimen was 
taken 16.xi.81 in Bataan). Using Semper's misidentified figure for comparison 
Fruhstorfer (1908) described hebetatrix as a wet-season form of schadenbergi, presum- 
ing Semper's less heavily brown shaded specimen to be the dry-season one. Female 
specimens of P. schadenbergi indeed vary considerably, particularly in the extent of 
brown colouration on the hindwing upperside and the size of the forewing white 
markings. Specimens have been seen in which the hindwing discal area is pure 
white and others in which brown dusting invades considerably into the discal area 
past the ocellar ring. The holotype of hebetatrix is intermediate between these 
extremes of variation. Data to hand are insufficient to correlate this variation 
with seasonal conditions. 

Material examined. 

Ptychandra schadenbergi, lectotype <$, Mindanao, coll. C. Semper 'Typus SMFL 
19' (SMN); 1 £ paralectotype, Mindanao: N. (SMN); 3 £ paralectotypes, Mindanao: 
N. (BMNH). Ptychandra schadenbergi f. hebetatrix, lectotype 9, Mindanao (BMNH). 

Mindanao: Baracata, Davao, 1 £, xii. 1922 (F. Grinnell) (BMNH); Calumbogan, 
2 J, 1 9, vii-viii. 1917 (CM); Davao, 1 <J, 1889 {Platen) (MNHU); Davao, 2 9 {Platen) 
(BMNH; Davao, 1 <J, 1 9, iii. 1890, (BMNH); Davao, 1 $, 1 J, 5.V.1962 (C. G. Tread- 
away) (BMNH); Sibulan, 1 <J, 3J.82 (BMNH); Sibulan, 1 9, i.xii.82, (SMN) ; Talaud, 
Malita, Davao, 1 J, 13.vi.61 {J. M. Jumalon) (BMNH); N. Mindanao, 2 £ (BMNH); 
N. Mindanao, 1 9 (SMN); 3 <J, 3 9 {Platen) (MNHU); N. Mindanao, 2 $, 1 9, 1903- 
1904 (/. Waterstradt) (BMNH); N. Mindanao, 2 <$, 3 9, (BMNH); N. Mindanao, 
2 9, 1920 (C. J. Grist) (BMNH). Luzon: i J, 1920 {E. C. Brabant) (BMNH). Pala- 
wan: 1 S (BMNH). 

Ptychandra mindorana Semper stat. rev. 

(Text-figs 9, 10; PL 2, fig. 5; PI. 3, fig. 10; PI. 5, figs 2i, 22) 

Ptychandra mindorana Semper, 1892 : 329. LECTOTYPE <$, Philippines: Mindoro (SMN), 

here designated [examined]. 
Ptychandra lorquinii mindorana Semper; Fruhstorfer, 1899 : 80; 1908 : 223; 191 1 : 330. 
Ptychandra lorquinii var. mindorana Semper; Gaede, 1931 : 320. 

Description. £. Forewing length 25-9-27-7 mm, n = 4. Fades. Forewing strongly 
convex distal of centre of costa. Scalloped appearance given to outer margins by long promi- 
nent white cilia interrupted at vein ends by dark brown ones. White chevrons and subapical 
fleck well marked. Forewing brand of type (b) (PI. 2, fig. 5) with pouched hair pencil contain- 
ing long mid-brown hairs. Pale elliptical source area for hair pencil elongate (3 mm x 1 mm). 
Hindwing margin rounded, evenly scalloped in cells CuA 1 , CuA 2 , M 2 , and M 3 . Tail at M 3 
not hooked and poorly developed. Underside dark brown streaked with deep brown and 
prominent metallic silvery blue lines in discal area. Full series of ocelli on hindwing, edged 
inwardly and marginally by diffuse pale metallic blue band and with those in cells M x and M 2 
confluent, that in cell M 2 inwardly displaced. Ocelli white-pupilled, ringed with black, light 



246 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

brown, buff and dark brown; the light brown and buff bands separated by a thin black line, the 
buff band being broadened distally. Antennae. Club tip rufous with base dark brown. Dorsal 
scaling on shaft grey, extending into club. Ventrally rufous with pale flecks also invading 
club. Genitalia. Tip of uncus notched ventrally (Text-fig. 9). 

$. Forewing length 27-4-30-8 mm, n = 4. Upperside forewing evenly convex along 
costa, of mid-brown ground colour with cream-coloured markings. Postdiscal bar extending 
from half-way along costa to M 2 and in line with a cream circular spot in cell M 2 above the 
submarginal ocellus. Off-white discal patch extending from half-way along the lower disco- 
cellular vein towards tornus but not crossing vein \A -\- iA. Fringes white interrupted with 
light brown at veins. Upperside hindwing discal area sandy brown becoming paler round 
costal ocellus. Hindwing margin rounded and evenly scalloped in cells iA + 2A, CuA x , 
CuA 2 , and M 3 . Tail at M 3 not hooked, bluntly terminated, short and narrow. Hindwing 
ocelli diffusely visible on upperside but with light brown rings in cells M 3 and CuA 1 prominent. 
Upperside cream areas reflected on underside. Forewing underside with three equal confluent 
ocelli in cells M v M 2 and M 3 ; that in cell M 3 inset. Hindwing cell Rs ocellus much enlarged, 
extending to M 2 and inset. Cell M 2 underside hindwing ocellus also inset and contiguous with 
that in cell M x . Ocellus white-pupilled ringed with brown, light brown, buff and brown. 
Antennae. Similar to male. 

Distribution. Mindoro. 

Biology. Said to fly in February (Semper, 1892). 

Discussion. Fruhstorfer (1911) treated mindorana as a subspecies of lorquinii 
without comment. However, the differences from lorquinii in secondary sexual 
characters and genitalia and wing shape lead us to consider it a separate species 
as originally described. The forewing brand closely resembles that of leucogyne 
except that the hair pencil has a more elliptical origin. 

Material examined. 

Ptychandra mindorana lectotype <$, Mindoro: Paluan, 23.il. 89 (SMN); 1 $, para- 
lectotype (SMN). 

Mindoro: Calapan, 1 <$, 1890-91 {Platen) (MNHU); Calapan, 1 <$, 15.V.1956 
(/. N. Jumalon) (BMNH); 2 $, 1 $ (BMNH); 2 $ {Platen) (MNHU); 3 J, 3 ? {Platen) 
(BMNH). 

Ptychandra leucogyne Felder & Felder stat. rev. 
(Text-figs 4, 13; PI. 1, fig. 2; PL 3, fig. 11; PI. 5, fig. 23, 24) 

Ptychandra leucogyne Felder & Felder, 1867 : 498; Fruhstorfer, 1899 : 81. LECTOTYPE 

$, 'Halmaheira' (BMNH), here designated [examined]. 
[Ptychandra schadenbergi Semper, 1886 : 59-60, pi. 11, fig. 7. Misidentification in part.] 
Ptychandra lorquinii leucogyne Felder & Felder; Fruhstorfer, 1908 : 223; 1911 : 330. 
Ptychandra lorquinii var. leucogyne Felder & Felder; Gaede, 1931 : 320. 

Description, g. Forewing length 24-2-25-1 mm, n = 4. Fades. Forewing costa weakly 
convex. Hindwing with slightly hooked tail on M 3 with acute not rounded end. Hindwing 
margin tending to be quadrate not circular. Upperside deep iridescent blue, darkened narrowly 
at margin. Cilia dark brown on forewing and light brown interrupted by dark brown at veins on 
hindwing. White subapical fleck and faint, pale blue antemarginal chevrons variably present 



REVISION OF GENUS PTYCHANDRA 



247 



on forewing. Forewing underside with four disjunct ocelli in cells Rs to M 3 , that in cell Rs 
sometimes reduced and that in cell M 3 inset. Brand and hair pencil, type (b) (PI. 1. fig. 2). 
Small black hair pencil in hindwing cell close to origin of Rs (PI. 5, fig. n). White origin of 
forewing black hair pencil, 2 mm x o-8 mm. Ocelli having white pupil ringed with black, 
light brown, buff, dark brown and fringed anteriorly and marginally with diffuse pale violet 
wavy line. Antennae. Club dark brown with rufous tip. Shaft with uniform grey-brown 
scales dorsally, rufous ventrally with paler flecks. Genitalia. Similar to those (Text-fig. 3) of 
schadenbergi, with serrated valve tips. Uncus with smooth unnotched tip. Gnathi smooth 
without dorsal teeth but narrowing distally and hooked. 

$. Forewing length 24 , 6-27-8mm, n = 4. Upperside predominantly off-white, dusted 
variably with brown basad. White forewing discal area extends to inner margin and hindwing 
white discal area extends to costa. Maximum convexity of forewing at about halfway along 
costa. Hindwing subquadrate with hooked tail at M 3 , terminated acutely. Forewing costal 
and apical areas dark brown, broken by subapical white fleck, two-thirds distad on costa and 
postdiscal white bar running from half way along costa towards white submarginal spot in 
cell M 2 . Faint brown ocelli in cells Rs and M 3 on forewing upperside. Upperside hindwing 
off-white with light brown submarginal area sometimes extending just beyond submarginal 
ocelli which are diffuse brown ringed with buff. Forewing underside with four discrete or 
almost discrete ocelli in cells Rs to M 3 , that in cell Rs sometimes reduced and cell M 3 inwardly 
displaced. Ocellus in cell M x , forewing, white-pupilled, ringed with brown, light brown, buff 
and brown, others, especially in M 2 , more broadly white. Antemarginal lines dark brown, 
yellow-brown and dark brown and cilia light brown. Antennae. Similar to those of male. 

Distribution. Luzon, Samar, Negros, Cebu, Halmaheira. This is the only 
Ptychandra recorded from Cebu. 

Biology. It has been observed (C. G. Treadaway, personal communication 
and specimens) that the two Ptychandra occurring on Luzon, lorquinii lorquinii 
and leucogyne, occupy different habitats. The more commonly taken species 
lorquinii lorquinii flies in secondary forest containing some primary areas while 
leucogyne, which is poorly represented in collections, occurs in glades in primary 
jungle. In the more commonly taken species males predominate but in leucogyne 
neither sex appears easily captured. This presumably reflects the more furtive 
habit of the latter species. 

Discussion- P. leucogyne has been confused with both lorquinii and schadenbergi. 
The males closely resemble lorquinii lorquinii and occur with this subspecies on 
Luzon. They are separated by habitat and distinguished easily by the number of 
forewing ocelli and the presence or absence of a hindwing brand. The hindwing 
tail and cell are significantly shorter and the forewing hair pencil is longer. The 
female was described as that of schadenbergi by Semper but has a wing and tail 
shape more characteristic of lorquinii. It may be conveniently distinguished 
from schadenbergi by the number of forewing ocelli and the extent of the white 
discal areas which in leucogyne reach to the inner margin of forewing and costa of 
the hindwing. Even strongly white-marked specimens of schadenbergi do not 
show this. 

The wing scales of this species and talboti have a tendency to be multitoothed. 
In the hindwing subcostal ocellus, the area intensively studied, four and five pointed 
scales are normal but three pointed scales predominate in lorquinii and schaden- 
bergi. 



248 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

The original description (Felder, 1867) is of a specimen said to be taken in Halma- 
heira by Lorquin, while Semper (1886) states that leucogyne flies on Batjan. How- 
ever, the facies of the lectotype is within the range of variation observed for leucogyne 
from Luzon and no other specimen of Ptychandra examined, except the lectotype 
of P. leucogyne, has been labelled as taken in the North Moluccas. Furthermore, 
Boisduval (1868) reports that Lorquin, the collector of the lectotype specimen, 
left San Francisco in 1857 to go to Luzon where he stayed for two years. He also 
took ship, apparently the 'No vara', from Manila later when he visited Celebes and 
Halmaheira. Luzon is a reasonable locality for this species and it is noteworthy 
that the Felders (1867) described several species (e.g. Ergolis luzonia, Charaxes 
amycus, Euripus clythia) from Luzon taken by Lorquin in the same publication 
as the description of leucogyne. It is thus possible that the lectot3^pe was taken 
in Luzon and not Halmaheira as described. Semper's record of the species from 
Batjan may have originated from the specimen actually of leucogyne but figured 
as $ schadenbergi (Semper, 1886) which was taken in Bataan, Luzon. 

Material examined. 

Ptychandra leucogyne, lectotype $, Halmaheira (Lorquin) (BMNH). 

Luzon: Bataan, 1 $, 16.xi.81, 'Paratypus SMFL 19b' (described as P. schadenbergi) 
(SMN); Los Banos, 1 $, vii. (B. P. Clarke) (USMN); Montalban, 1 $, 30.iv.19n 
(A. E. Wileman) (BMNH); Mt Makiling, 1500 ft, 8 $, 20. xi. 1955, 3.viii.i956, 31.VUL 
1966, 4.IX.1966 (C. G. Treadaway) (BMNH); Paete, 1 $, vi. (B. P. Clarke) (USNM); 
Tarlac 1 $ (Browne) (BMNH); 1 $ (BMNH). Cebu: Minglanilla, camp 7100ft, 
1 <$, 1. x. 1958, woodland (/. N. Jumalon) (BMNH). Negros: i ^, lowlands, ii. 96 
(/. Whitehead) (BMNH). Samar: Antiago Valley, 1 <J (/. /. Mounsey) (UM). 



Ptychandra talboti Hobby stat. n. 
(Text-figs 5a, b; PL 1, fig. 1; PL 3, fig. 9; PL 5, figs 25, 26) 

Ptychandra schadenbergi talboti Hobby, 1940 : 220, pi. 4, figs 1, 2. Holotype °-. Borneo: 
Sarawak, Mt Dulit (BMNH) [examined]. 

Description. <-£. Forewing length 19-1-20-8 mm, n = 4. Facies. Upperside deep iri- 
descent blue-purple with narrow darkened margins. Cilia short and white, interrupted by 
brown at veins, giving scalloped appearance. Forewing costa strongly convex. Forewing 
brand and hair pencil of type (a) (PI. 1, fig. 1). Hair pouch below CuA 1 forewing absent. 
Hindwing quadrate, strongly excavate in cell M 2 and weakly in cells M 3 , CuA 1 and CuA 2 - 
Tail at M 3 weakly hooked and obtusely terminated. Hindwing with prominent black hair 
pencil (PI. 3, fig. 9) arising in cell at base of Rs and lying across cell. Underside forewing with 
four submarginal ocelli in cells Rs-M 3 , with those in cells Rs and M 3 slightly reduced. Costal 
ocellus of hindwing only slightly larger than remaining ocelli but inwardly displaced. Ocelli 
discrete with blue-white pupil ringed with black, mid-brown and dark brown and fringed 
incompletely, interiorly and marginally with a faint mauve line. Marginal lines dark brown, 
light brown, dark brown and faint mauve. Antennae. Club slightly flattened, rufous, darker 
dorsally. Shaft black-scaled, with ventral white flecks basal to each segment. Genitalia 
(Text-fig. 5a). Gnathi toothless and weakly hooked at tip. Uncus smooth and rounded 
distally. Valvae without strongly sclerotized distal processes but with weakly toothed and 
serrated tips. Aedeagus (Text-fig. 5b) poorly developed, with only small areas of sclerotization. 



REVISION OF GENUS PTYCHANDRA 249 

§. Forewing length 21 -i mm. Fades. Wing shape similar to male. Upperside forevving 
ground colour chocolate brown with white costal bar divided into three by darkened veins 
and extending to M z . Subapical white fleck absent or vestigial. Discal white patch not 
extending to tornus but pointed distally. Submarginal white chevrons absent. Fringes 
dark brown. Hindwing discal area white, extending to costa and submarginal ocelli. Ocelli 
visible with eye spots of those in cells M 3 and CuA 1 sharply defined. Fringes greyish inter- 
rupted by smokey brown. Underside forewing costal band extending to M 3 . Discal white 
patch as upperside. Four subequal submarginal ocelli in cells Rs to M 3 in band sloping to 
tornus, fringed inwardly and marginally by a violet-tinged white band. Hindwing with full 
series of well defined ocelli with that in cell Rs enlarged and displaced inwardly. Discal area 
white, traversed by three much interrupted dark brown bands. Ocelli discrete, with white 
pupil ringed successively with dark brown, buff and mid-brown and externally fringed with 
diffuse white line. Antennae. Uniformly rufous except club darkened dorsally. Club slightly 
flattened. 

Distribution. Borneo. 

Biology. Taken in or close to primary montane forest. 

Discussion. Only three female specimens ascribable to this species were ex- 
amined. The female holotype from Mt Dulit was meticulously described by Hobby 
(1940). The specimens from Mt Kinabalu were considerably less brown dusted 
than the former, the brown marginal shading extending into the discal area beyond 
the submarginal ocelli in contrast to that of the Mt Dulit specimen in which it 
extends to halfway inward from the margin towards the submarginal ocelli. The 
Mt Kinabalu specimens also differed by having much paler discal areas on the 
underside with reduced brown banding and a slightly different hindwing shape, 
that of the holotype being more rounded with a less hooked tail and similar to that 
of schadenbergi. These differences are as extreme as the variation found within 
the series of female schadenbergi or leucogyne examined, but it may be that diver- 
gence to at least subspecific level has occurred on the Bornean mountains which 
will be revealed by examination of a longer series. 

This species is considerably smaller than the Philippine Ptychandra and is thus 
distinguished from these biometrically by many of the parameters used. It super- 
ficially resembles leucogyne, particularly in general facies and possession of four 
forewing ocelli. It can be distinguished in the male by its size and secondary 
sexual characters, particularly the lack of the hair pouch below CuA v Despite 
the small size of this species, the forewing hair pencil is exceeded in length only 
by that of mindorana. In the female the white discal areas never reach the inner 
margin or costa, being margined by a brown area. In both sexes the forewing 
costa is markedly convex, not almost straight as in leucogyne. It is the only Pty- 
chandra species to have four not five rings to the hindwing ocelli. The genitalia, 
particularly the aedeagus, differ considerably from those of other members of the 
genus. They are more delicate, less spiny structures and the aedeagus is reduced 
to a poorly defined chitinous fold. 

Material examined. 

Ptychandra schadenbergi talboti, holotype $, Sarawak: 4600 ft, 17.x. 32 (B. M. 
Hobby & A. W. Moore), moss forest, Oxford University Expedition (BMNH). 



250 H. J. BANKS, J. D. HOLLOWAY & H. S. BARLOW 

Borneo: Mt Kinabalu, between 5500-6000 ft, 3 $, 1 $, 25.vii, 29.vii, 2.viii, 3.ix, 
1965 (H. J. Banks, H. S. Barlow & J. D. Holloway) (BMNH) ; Mt Kinabalu, 5500 ft, 
1 cJ, 1 $, 22, 24.iii.1974 (Shoichi Iwanaga) (Iwanaga coll., Osaka, Japan) (examined 
from colour photographs); Mt Trus Madi, 7800 ft, 1 3, 3.viii.i956 (BMNH). 



Ptychandra negrosensis sp. n. 

(PI. 2; fig. 6; PI. 3, fig. 12; PL 5, figs 27, 28) 

Description, g. Forewing length 28-7111111. Fades. Forewing slightly falcate with 
maximum convexity two-thirds distad on costa. Upperside deep iridescent blue with promi- 
nent broad rhomboidal white apical fleck, diffuse white antemarginal chevrons on forewing, 
appearing as pale blue wedge, broader apically, and white diffuse antemarginal line in hindwing. 
Small, purple ocellus in cell CuA 2 . Forewing hair pencil and brand structure type (c) (PI. 2, 
fig. 6) with pouched hair pencil and white mealy scales in brand. Hindwing quadrate, prominently 
tailed at M 3 with excavation of cell M 2 much greater than of cell M 3 , producing a strong hook. 
Tail terminated bluntly. Prominent black hair pencil (4-6 mm) in cell at origin of Rs (PI. 3, 
fig. 12). Underside forewing dark brown with lighter brown transverse wavy lines inwardly, 
ocelli in cells M 1 -M 3 , that in il/ x reduced. Antemarginal chevrons as upperside in shape, 
violet-tinged white, well-marked with prominent apical fleck. Slight metallic marbling in 
underside discal area. Full series of hindwing submarginal ocelli, with cell Rs ocellus inwardly 
displaced and much enlarged. Ocelli in cells M x and M. 2 contiguous, others discrete; white 
pupilled ringed with mid-brown, light brown, buff and mid-brown lines and with diffuse violet 
band fringing ocelli, inward and marginally. Antennae. Club not well defined but with dark 
brown tip. Shaft grey dorsally, rufous exteriorly and paler ventrally. Genitalia. Gnathos 
smoothly curved distally with small dorsal spines extending about one-third of length. Uncus 
unnotched at end. Valves with two heavily toothed terminal processes similar to Text-fig. 2 
but without middle process. 

9. Forewing, length 34-3 mm, not falcate. Fades. Upperside forewing brown with broad, 
white, entire subapical band running from half way along costa to anal angle. Subapical white 
fleck present. Hindwing prominently tailed at M 3 as in male. Upperside hindwing brown 
with dull brick red discal area. Two forewing ocelli in cells M x and M 2 , almost conjoined. 
Submarginal ocelli visible on upperside hindwing, particularly cells M 2 and M 3 , ringed with 
rusty brown and white pupilled. Underside discal area whitish with brown transverse wavy 
lines. White patch at costa distad of cell Rs ocellus, which is enlarged and displaced inwardly. 
Ocelli in hindwing cells ft to 1^ + 2 A with white pupil ringed with mid-brown, light brown, 
buff and mid-brown lines and visible on upperside. Ocellus in cell M 3 displaced outwardly 
those in cells M x and M 2 contiguous. 

Distribution. Negros. 

Biology. This species was taken in rain forest in a thick undergrowth of ferns 
and vines on an overcast day. It was feeding from cut grass stems in the company 
of amathusiids and Neptis species (/. TV. Jumalon, personal communication). 

Discussion. This new species is easily distinguished from other Ptychandra 
by its large size. In the two specimens known the forewing ocellar band is reduced 
to two or one spot only. The prominent hindwing band distinguishes the male 
from other species while the female can be told by the very broad unbroken forewing 
subapical white band and the reddish discal area of the hindwing upperside. The 



REVISION OF GENUS PTYCHANDRA 251 

cJ genitalia are similar to those of southern lorquinii and without a series for com- 
parison cannot be confidently distinguished. 

Material examined. 

Holotype <$, Negros: Bayawan, x. 1963 (J. N. Jumalon) (BMNH). 
Paratype (allotype) <j>, Negros: Bayawan, x. 1963 (/. N. Jumalon) (BMNH). 



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xiv + 380 pp., 2+49 pis. Wiesbaden. 
Staudinger, O. 1884-1888. Exotische Tagfalter. In Staudinger, O. & Schatz, E., Exotische 

Schmetterlinge 1 (1) : vi + 333 PPJ (2) : iv + 100 pis, 1 map. Fiirth. 



INDEX 

Synonyms and unavailable names are in italics ; principal references are in bold. 

alitha, 224 
amycus, 248 



bazilana, 221, 242 

caerulans , 238, 239, 242 
clythia, 248 

Elymniinae, 224 
europa, 224 

hebetatrix, 244 

Lethe, 221, 224 

Lethini, 224 

leucogyne, 220, 221, 225-234, 246 

leytensis, 221, 241 

lorquini, 237 

lorquinii, 218, 220, 221, 225-234, 237, 239, 

243 
luzonia, 248 

Mandarinia, 224 
Melanitini, 224 



mindanaensis , 241 
mindorana, 221, 245 
minerva, 224 
Mycalesini, 224 
Mycalesis, 224 

negrosensis, 220, 250 
Neptis, 220, 250 
Neorina, 224 

obscurior, 239 
Orsotriaena, 224 

perseus, 224 

plateni, 221, 232-234, 241 

Ptychandra, 218, 222, 224, 236 

schadenbergi, 218, 220, 221, 225-234, 244, 
246 

talboti, 217, 220, 221, 231, 248 

Zemeros, 221 



H. J. Banks, M.A., Ph.D., A.R.I.C. 

Division of Entomology 

C.S.I. R.O. 

P.O. Box 1700 

Canberra, A.C.T. 2601 

Australia 



J. D. Holloway, M.A., PhD. 

TlLLINGLEA 
TlLLINGBOURNE PARK 
WOTTON 

Dorking 
Surrey RH5 6QL 

H. S. Barlow, M.A. 
P.O. Box 295 
Kuala Lumpur 
Malaysia 



PLATE i 

Forewing male brand structures in Ptychandra 
Fig. i. Type (a), talboti. 
Fig. 2. Type (b), leucogyne. 
Fig. 3. Type (c), lorqninii lovquinii. 



Bull. Br. Mil-,, not. Hist. (Ent.) 32, 6 



PLATE 1 




PLATE 2 

Forewing male brand structures in Ptychandra 
Fig. 4 schadenbergi. 
Fig. 5. Type (b), mindorana. 
Fig. 6. Type (c), negrosensis. 



Bull. Br. Mas. nut. Hist. (Ent.) 32, 6 



PLATE 2 




6 *-•.-*• 



PLATE 3 



Hindwing male brand structures in Ptychandva 



Fig. 7 
Fig. 8 
Fig. 9 
Fig. io 
Fig. ii 
Fig. 12 



lorquinii lovquinii. 

schadenbergi. 

talboti. 

mindorana. 

leucogyne. 

neerosensis. 



Bull. Br. Mas. mil. Hist. (Ent.) 32, 6 



PLATE 3 





■ 

■•.-..- 


■ T • 


-''' 


'..: 




, 


7 


.."■' 







PLATE 4 



Fig. 


13- 


Fig. 


14. 


Fig. 


15- 


Fig. 


16 


Fig. 


17- 


Fig. 


18 


Fig. 


19 


Fig. 


20 



Ptychandra lorquinii lorquinii, lectotype £. 

P. lorquinii lorquinii, £• 

P. lorquinii plateni, <£. 

P. lorquinii bazilana, 9- 

P. lorquinii leytensis, $. 

P. lorquinii, £, Bohol. 

P. schadenbergi, <$. 

P. schadenbergi, §. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 6 



PLATE 4 










PLATE 5 


Fig. 


21 


Ptychandra mindorana, <j>. 


Fig. 


22 


P. mindorana, $. 


Fig. 


2 3 


P. leucogyne, g. 


Fig. 


2 4 


P. leucogyne, lectotype $■ 


Fig. 


25 


P. /a/60/i, (J. 


Fig. 


26 


P. teftoft', $, Mt Kinabalu. 


Fig. 


27 


P. negrosensis, holotype <J. 


Fig. 


28 


P. negrosensis, allotype 5- 



Bull. Hi. Mus. nat. His/. (Ent.) 32, 6 



PLATE 5 





ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera) . Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 

i9 6 5- £3-25- 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae) . Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae) . Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea) . Pp. 908: 686 text-figures. November, 1969. 

£*9- 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. 
February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text -figures. December, 1973. £9.55. 

printed by Unwin Brothers Limited the gresham press old wokinc surrey England 



0» I T-- 



A TAXONOMIC REVISION OF THE 
TINISSINAE OF THE WORLD 
(LEPIDOPTERA : TINEIDA: 




G. S. ROBINSON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 7 

LONDON : 1976 



A TAXONOMIC REVISION 

OF THE TINISSINAE OF THE WORLD 

(LEPIDOPTERA : TINEIDAE) 



BY 

GADEN SUTHERLAND ROBINSON 

i. 



Pp. 253-300; 16 Plates; 10 Text-figures 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 7 

LONDON : 1976 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 7 of the Entomology 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. not. Hist. (Ent. 

ISSN 0524-6431 



Trustees of the British Museum (Natural History), 1976 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 25 March, 1976 Price £6.25 



A TAXONOMIC REVISION 

OF THE TINISSINAE OF THE WORLD 

(LEPIDOPTERA : TINEIDAE) 

By G. S. ROBINSON 



CONTENTS 

Synopsis 

Introduction 

Acknowledgements 

Abbreviations 

Checklist of Tinissinae 

Geographical distribution 

Biology 

Phylogeny 

Morphology . 

Tinissinae Gozmany & Vari 

Key to male Tinissinae based on genitalia 
Key to female Tinissinae based on genitalia 
Tinissa Walker ...... 

Leptozancla Meyrick ..... 

References ....... 

Index ........ 



Page 

255 
255 
256 
256 
256 
257 
257 
257 
259 
261 
263 
264 
266 
297 
299 
300 



SYNOPSIS 
The subfamily Tinissinae (Lepidoptera) and its constituent genera Tinissa Walker and 
Leptozancla Meyrick are redescribed and redefined. Thirty-four species are described, sixteen 
of them new; one new subspecies is described and two new synonyms are established. Two 
species are transferred to Tinissa from other genera. Keys to the species are given and the 
possible phylogeny of the subfamily is discussed. 

INTRODUCTION 

The subfamily Tinissinae was erected by Gozmany & Vari (1973). It contains 
two genera, Tinissa Walker and Leptozancla Meyrick, with thirty-two and two 
species respectively. The group is exceptional within the Tineidae in that the 
juxta of the male is usually extensively modified and usurps the function of the 
valves which are actually lost in Leptozancla. This feature was not noticed by 
Diakonoff (1967) or Gozmany & Vari (1973). 

All primary types except three have been examined; in these three cases para- 
lectotypes were available or the primary type had been adequately described and 
illustrated. 

Size measurements given are double the distance from the wing apex to the 
centre of the thorax. 

The terminology used in descriptions of male and female genitalia follows Klots 
(1956). 



256 



G. S. ROBINSON 



Genitalia dissections were made and subsequently mounted in conventional 
fashion; in this group, however, it is necessary to separate the uncus from the 
valve-juxta complex in order to view it from the ventral side. Male genitalia 
dissections therefore involved cutting the vinculum laterally and severing the 
membrane anterior to the tip of the subscaphium and 'unrolling' the genitalia; 
the uncus is thus viewed from the ventral side and the valve-juxta complex and 
saccus from the dorsal side. Chlorazol Black E was used for staining preparations 
and Euparal was employed as a mounting medium. 

ACKNOWLEDGEMENTS 

I am most grateful to Dr D. R. Davis, National Museum of Natural History, 
Washington and to Dr A. Diakonoff, Rijksmuseum van Natuurlijke Historie, 
Leiden, for the loan of specimens. 

I am indebted to my colleagues, Dr K. Sattler and Mr P. E. S. Whalley, for their 
comments and advice during the preparation of this paper. Photographs were 
taken by the Photographic Section, BMNH. Mr B. S. Martin of the Electron 
Microscope Unit, BMNH, supervised my taking the Stereoscan electronmicrographs. 

ABBREVIATIONS 

BMNH British Museum (Natural History). 

MAK Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn. 

MNHN Museum National d'Histoire Naturelle, Paris. 

RNH Rijksmuseum van Natuurlijke Historie, Leiden. 

TM Transvaal Museum, Pretoria. 

USNM National Museum of Natural History, Smithsonian Institution, Washington. 



CHECKLIST 
TINISSA Walker 

POLYMNESTRA Meyrick 
albipuncta sp. n. 
amboinensis sp. n. 
araucariae sp. n. 
bakeri sp. n. 
baliomicta Meyrick 
chalcites sp. n. 
chaotica sp. n. 
cinerascens Meyrick 
convoluta sp. n. 

cultellata (Gozmany & Vari) comb. n. 
distract a Meyrick 
dohertyi sp. n. 
errantia sp. n. 
eumetrota Meyrick 
goliath sp. n. 
indica sp. n. 
insignis Zagulajev 
insular ia sp. n. 
kidukaroka sp. n. 
krakatoa sp. n. 



OF TINISSINAE 

palmodes Meyrick 
parallela sp. n. 
philippinensis sp. n. 
phrictodes Meyrick 
poliophasma Bradley 
polysema Zagulajev 
*polystacta (Meyrick) 

\perilithias (Meyrick) 
rigida Meyrick 

chloroplocama Meyrick syn. n. 

heterograpta Meyrick syn. n. 
ruwenzorica Gozmany 
spaniastra Meyrick 
torvella torvella Walker 
torvella mysorensis subsp. n. 
transversella (Walker) comb. n. 
LEPTOZANCLA Meyrick 

PHILAGRIAS Meyrick 
*talaroscia Meyrick 
zelotica (Meyrick) 

*paralectotype only examined 
f holotype not examined 



REVISION OF THE TINISSINAE 257 

GEOGRAPHICAL DISTRIBUTION 

The subfamily Tinissinae contains exclusively Old World taxa. The genus 
Leptozancla is Ethiopian; its two species are known from montane localities in 
Kenya and Ethiopia. Tinissa contains five Ethiopian species; four of these are 
known from montane localities in Uganda and Ethiopia and one is only known 
from South Africa. The remaining twenty-seven Tinissa species are Indo-Aus- 
tralasian, having been collected within a zone, the western limits of which are Sri 
Lanka (Ceylon), south India and Sikkim, the eastern limits Taiwan, the Philippines 
and the Solomon Islands, and the southern limit Queensland. Several species 
have extensive distributions (e.g. Tinissa insularia, T. rigida) but many (e.g. 
T. krakatoa, T. chaotica) are known only from a single locality. It is likely that 
most species are under-collected. Of the 211 specimens of Tinissa known to me, 
representing 33 taxa, seven species account for 155 specimens while 15 taxa are 
represented by a single specimen. It is, therefore, likely that there are many 
more species of Tinissinae to be discovered and that present distributional records 
are incomplete. From the imperfect records of distribution presently available, 
there is a concentration of species in the Papuan region. Only five species are 
known from the Asian mainland. Nine species are recorded from New Guinea 
and fifteen from Buru to the Solomon Islands. 

BIOLOGY 

All Tinissine species, with the exception of Leptozancla talaroscia and Tinissa 
polystacta, have been collected in tropical rain forest and their distributions seem 
to be restricted to this habitat type within about 20 north and south of the equator. 
No published host records exist for this group, but Meyrick, in a manuscript note- 
book preserved in the Microlepidoptera Section, BMNH, records (notebook no. 
12, page 58) Tinissa torvella 'on fungus on bamboo'. Mey rick's collection contained 
T. torvella, T. indica and T. insularia determined as 'torvella'' so it is uncertain 
as to which species this record really relates. It is enough to suggest, however, 
that the group is fungivorous. Specimens of Tinissa have been collected in every 
month of the year both north and south of the equator and the species probably 
breed continuously. Specimens are generally rare (see comments in 'Geographical 
Distribution') . 

PHYLOGENY 

No fossil evidence is available to strengthen any phylogenetic assumptions made 
about this group and only morphological and geographical evidence is here taken 
into account. I consider the Tinissinae to have originated in Africa where the 
remaining species are all markedly dissimilar and where all the species with a 
5-segmented maxillary palp are found. Reduction of the number of maxillary 
palp segments is considered to have been a single and early occurrence in the history 
of the group, antedating the migration of an early Tinissa stock to Asia. Tinissa 
phrictodes is rather different from the other Asian species; the structure of the 
uncus is similar to that of T. polystacta, suggesting that it is an early derivative 
from the first 'African-type' immigrant stock. 



258 



G. S. ROBINSON 




krakatoa 

cinerascens 

distracta 

amboinensis 

errantia 

chaotica 

convoluta 

palmodes 

araucariae 

chalcites 

albipuncta 

eumetrota 

polysema 

philippinensis 

rigida 

baliomicta 

parallela 

indica 

transversella 

dohertyi 

bakeri 

insularia 

goliath 

kidukaroka 

insignis 

phrictodes 

poliophasma 

spaniastra 

ruwenzorica 

torvella 

polystacta 

cultellata 

talaroscia 
zelotica 



Fig. i. Suggested phylogeny of the Tinissinae. 



REVISION OF THE TINISSINAE 



259 



The occurrence of 'pouches' in the female eighth tergite, while a useful diagnostic 
character, seems to be of little phylogenetic significance. Species with pouches 
show little similarity to each other when other characteristics are considered and 
the structure of the pouches suggests that they have arisen independently on 
several occasions. Two major groups of Asian species are recognized. The first 
group is of four quite closely related species (T. cinerascens , distracta, krakatoa, 
amboinensis) , remarkable for the narrowness of the male saccus, which are grouped 
with a series of ten species of progressively divergent genital morphology but 
generally with a pale wing pattern. This group is probably monophyletic and 
contrasts with the darker-winged second group of eleven species which form three 
quite well-defined species-groups. I consider Tinissa torvella to be a remnant of 
a separate invasion of Asia of a line otherwise represented by T. ruwenzorica. I 
consider Leptozancla to be monophyletic from the earliest Tinissine stock; its 
two species are widely divergent. Text-fig. 1 illustrates my concept of the phylo- 
genetic relationships of the Tinissinae. The affinities of the subfamily remain 
obscure and I am, at this stage, content to think of the Tinissinae as having arisen 
from a generalized Tineoid-Psychoid stock. 

MORPHOLOGY 

The morphology of the Tinissinae is described in detail below but comment is 
required on the peculiar structure of the male genitalia. Text-fig. 2 is a schematic 
diagram of the male genital structures of a Tinissine. Features which should 
be noted are the fusion of an enormous juxta with the valve and the presence of 
a process which arises from the membrane separating the internal surfaces of the 
valve and juxta. The labis should be noted, a posterior extension of the transtilla, 
usually close to or in contact with the aedeagus. The characteristic form of the 
uncus and subscaphium are also shown. 

Earlier authors have had different interpretations of male Tinissine genitalia; 
the following table equates the terminology used. 



Diakonoff (1967) 


Gozmany & Vari (1973) 


Present work 


valva 


ventral lobe of valve; 
hairy clavate appendage 


juxta 


anellus lobe ; triangular 
hairy process 


dorsal lobe of valve 


process arising from mem- 
brane between valve and 
juxta 


transtilla; slender process 
directed rostrad 


transtillae 


apodeme of valve 


elevated bristly knob 


anellus with two lobate 
arms 


valve 


— 


serrate cornuti 


spicules on outer surface of 
aedeagus 


tegumen 


uncus 


uncus 



260 



G. S. ROBINSON 



It is apparent that the valve-juxta complex of the Tinissinae has not been noted 
by earlier authors who have interpreted the complex as being wholly composed 
of the valves. In most species, a membranous line clearly separates the valves 
and the juxta and there is no evidence of a medio ventral division of the sclerotized 
portions of the complex as would be expected if the complex were composed of 
ventrally extended valves. In several species (e.g. Tinissa torvella), the juxta 
is reduced and not closely fused with the valves, but in several species the degree 
of fusion is such that it is difficult to trace the margin of the juxta. 

In the female genitalia of Tinissa spaniastra, Gozmany & Vari's (1973) 'minute 
phylliform signum' is a foreign body, probably a scale. 



uncus 



process from membrane 
between valve and 
juxta 




carina 



-juxta 



saccus 



aedeagus 



Fig. 2. Schematic diagram of male genitalia of a Tinissine. 



REVISION OF THE TINISSINAE 261 

TINISSINAE Gozmany & Vari 

Tinissinae Gozmany & Vari, 1973, Transv. Mus. Mem. 18 : 84. Type-genus: Tinissa Walker, 
1864. 

Frons loosely covered with short, pale scales, predominantly forward-directed, convergent 
in midline of head. Mandible present (presence uncertain in Leptozancla) , thinly sclerotized, 
four times as long as broad at base. Proboscis almost as long as second segment of labial 
palpus, the two galeae only loosely associated, if at all; epipharynx small, triangular, protu- 
berant (proboscis absent and epipharynx reduced in Tinissa polystacta). Maxillary palpus 
with five short segments, folded [Leptozancla, Tinissa polystacta, T. cultellata) , or three segments, 
first short, second twice as long, third 1-5 times as long as second. Three-segmented maxillary 
palpus with conspicuous tuft of divergent pale scales from terminal segment. Labial palpus 
with three segments, second and third approximately same length, first about one-third length 
of second; terminal segment upturned to about 6o c , second segment with ventral tuft of elon- 
gate scales projecting anteriorly and with lateral bristles. Antenna extending to about one- 
half of fore wing costa in Tinissa, almost to or to apex in Leptozancla, £ with elongate cilia 
(PI. 1, fig. 1), $ cilia shorter (PI. 1, fig. 2) (cilia very long in Tinissa chaotica, reduced in length 
or absent in Leptozancla zelotica, Tinissa ruwenzorica, T. poliophasma, T. spaniastra; cilia 
short but dense in Leptozancla talaroscia). Scape with stiff downward-pointing pecten. Fore 
wing (Text-fig. 3) brownish, with or without pale (usually white or cream) spots and/or trans- 
verse striae, somewhat rectangular (narrow, apex oblique in Leptozancla talaroscia), 5-15 mm 
in length; M present or absent in cell; R h to apex or just on to termen; 7? 4 and R h separate, 
stalked or completely fused. With (Tinissa) or without (Leptozancla) elongate oval patch 
(PI. 1, fig. 3) of small, flat, ovate scales (PI. 1, fig. 4) on ventral surface of wing between A 1+2 
and posterior margin. Hind wing with patch of rough, pale scales opposite fore wing patch, 
anterior to Sc + i? x (absent in Leptozancla). Venation usually complete but M sometimes 
absent from cell. Female frenulum with two or three spines; if three, one may be shorter 
than other two (§ of Leptozancla unknown). Hind tibia with conspicuous distal hair tuft 
or with smooth broad scales, first hind tarsal segment smooth-scaled (with rough, raised scaling 
in Tinissa spaniastra) ; mid tibia with two oblique brownish transverse bands (absent in Tinissa 
cultellata) on outer face. 

Genitalia $. Simple, pouch-shaped corema (PI. 7, fig. 47) present in tergosternal membrane 
(absent in Tinissa polystacta, T. phrictodes and T. chaotica). Eighth segment well sclerotized, 
sternite with pair of lateral processes in Tinissa torvella and T. ruwenzorica. Saccus always 
large and conspicuous though sometimes narrow; vinculum extending dorsally to form an 
almost complete ring, tips separated by membrane. Tegumen and gnathos absent. Uncus 
completely divided, a pair of lobes attached to vinculum and each other by membrane (fused 
with vinculum in Leptozancla zelotica, Tinissa cultellata and T. polystacta) ; shape of lobes and 
presence or absence of processes or invaginations highly diagnostic at species level. Juxta 
large, closely appressed to valves, diversely modified and in many species taking over function 
of valves. Valves usually reduced, not as diversely shaped as juxta, absent in Leptozancla. 
Variously shaped and modified process arising from membrane between valve and juxta present 
in most Tinissa species, absent in Leptozancla. Labides present, usually a pair of lobes, some- 
times fused, dorsal to aedeagus, or strongly developed, forming posteriorly directed spines 
(Leptozancla) ; in several species pair of labides lobes ventral to aedeagus ; labis absent in a 
few species. Subscaphium elongate, extending from edge of anal orifice to proximity of base 
of valves, usually narrow, sclerotized, conspicuous; in two species subscaphium broadened 
and laterally evaginated to form a pair of posteriorly directed spined processes. Aedeagus 
of various shapes and sizes, with or without ventral carina near apex, cornuti absent; one 
species (Tinissa spaniastra) with a line of fine spicules on outer surface of aedeagus. 

Genitalia 9- Seventh tergite with posterior medial lobe in several species. Eighth sternite 
variously modified with posterior and anterior emargination, keel-shaped in several species, 
often setose, with or without conspicuous setae at posterior margin. Ostium similarly diverse, 



262 



G. S. ROBINSON 



at various angles to plane of eighth sternite, in several species with marked emargination of 
ventral lip. Antrum always present, sclerotized, often divided into two sections by membra- 
nous zone at point of junction with ductus seminalis although in several species sclerotization 
is continuous. Ductus bursae thin-walled but strengthened by regular transverse constrictions 
which may, additionally, be rugose with fine superficial pimpling; ductus bursae anteriorly 
with a single loop to the right. Corpus bursae with extremely thin walls, without signum. 
Eighth tergite variously modified; with or without posterior medial emargination; with or 
without various pouch formations in tergite wall or at anterior margin; with transverse ridging 
in T. cultellata; more or less setose, a row of setae of varying degrees of conspicuousness at 
posterior margin; with or without evagination of ventral surface to form lateral flaps around 
base of ovipositor. Ovipositor elongate, apophyses posteriores almost reaching tips of apophy- 
ses anteriores when ovipositor extruded; lightly sclerotized region anterior to anal papillae 
extending to posterior fold of ovipositor with fine transverse ridging on ventral surface. 




A l+2 



CuP 



Fig. 3. Venation of Tinissa torvella torvella Walker, $, Sri Lanka. 



REVISION OF THE TINISSINAE 263 

Remarks. Intraspecific variation in species represented by long series is quite 
limited within this group but specimens may be variable in size (e.g. Tinissa insu- 
laria); females are generally larger than males. Venation is variable: R 4 and R 5 
in the fore wing may be separate, stalked or fused in Tinissa torvella. The number 
of spines of the female frenulum may be two or three in the same species; one of 
three spines may be shortened. Specimens of T. torvella from Sri Lanka (Ceylon) 
collected in 1970 have the fore wings blackish brown with a purplish sheen. Older 
material is distinctly faded, a dull medium brown. It is likely that all older materia) 
described in this paper is faded to some extent. Aberrant specimens have been 
encountered; the holotype of T. palmodes has asymmetric hind wing venation and 
a specimen of T. baliomicta has deformed genitalia and abdomen (genitalia slide 
no. W 02). A female specimen of T. insularia has deformed genitalia and possibly 
exhibits gynandromorph characteristics in the elongation of the left side of the 
seventh segment. In several species there is distinct correlation between the shape 
and size of the aedeagus and of the ostium and antrum. An emargination of the 
ventral lip of the ostium accommodates the carina of the aedeagus. The patch 
of raised scales anterior to Sc -\- K x on the hind wing is often worn off or obscured 
by the fore wing. Gozmany & Vari (1973 : 84) suggest that the female lacks a 
corethrogyne, a dense tuft of scales from the seventh segment surrounding the 
ovipositor, but I have found a corethrogyne in all species examined. The apophyses 
anteriores apparently arise from the eighth sternite in this group. 



Key to male Tinissinae based on genitalia 

Males of polysema, philippinensis, insignis, avancariae, palmodes, eumetrota, convoluta, krakatoa. 
tr an sver sella, dohertyi, goliath, kidukaroka and poliophasma are not known. 

1 Valve present, apodeme visible (TINISSA) ....... 3 

- Valve absent, apodeme not visible (LEPTOZANCLA) 2 

2 Labides two posteriorly directed spines, juxta with distinct ventro-medial suture; 

aedeagus more than i-i mm long (PI. 11, fig. 66) .... zelotica (p. 298) 

- Labides four posteriorly directed spines, juxta without ventro-medial suture; 

aedeagus less than 0-7 mm long (PI. 10, fig. 65) . . . talaroscia (p. 297) 

3 Eighth sternite with posterior process at each corner (PI. 10, figs 61, 63) . . 4 

- Eighth sternite without posterior process at each corner ..... 5 

4 Lobes of uncus with small digitate lateral process; aedeagus 0-9 mm long (PI. 10, 

fig. 60) .......... ruwenzorica (p. 292) 

- Lobes of uncus without small digitate lateral process; aedeagus 1 -6 mm long (PI. 10, 

fig. 62) . . . . . . . . . . . torvella (p. 292) 

5 Coremata present in tergosternal membrane of eighth segment .... 8 

- Coremata absent from tergosternal membrane of eighth segment .... 6 

6 Tips of uncus lobes bifid, terminating in pair of large, thorn-like spines (PI. 8, fig. 50) 

chaotica (p. 271) 

- Tips of uncus lobes not bifid .......... 7 

7 Transtilla cowl-shaped, extended posteriorly; subscaphium rod-like anteriorly; 

uncus fused with vinculum; aedeagus small (o-8o x 0-06 mm) (African species) 

polystacta (p. 294) 



264 G - s - ROBINSON 

- Transtilla not developed; subscaphium broad, blade-like anteriorly; uncus not fused 

with vinculum; aedeagus large (i-io x 0-14 mm) (Australasian species) (PI. 9, 

fig. 58) ......... . phrictodes (p. 289) 

8 Aedeagus with line of fine spines or spicules on external surface (PI. 9, fig. 59) 

spaniastra (p. 291) 

- Aedeagus smooth, or with only one or two thorn-like carinae on external surface . 9 

9 Uncus fused with vinculum (PI. 10, fig. 64) .... .cultellata (p. 296) 

- Uncus separated from vinculum by narrow membranous zone . . . . 10 

10 Subscaphium evaginated to form pair of large, spined, digitate processes . . 11 

- Subscaphium without processes . . . . . . . . .12 

11 Aedeagus 1 -8 mm long, with large, conspicuous carina from three-quarters, almost 

reaching tip (PI. 9, fig. 55) ........ indica (p. 282) 

- Aedeagus without carina, i-i mm long (PI. 9, fig. 56) .... bakeri (p. 284) 

12 Anterior portion of saccus slender, parallel-sided and elongate, at least seven times 

as long as broad ........... 13 

- Saccus triangular or, if parallel-sided, less than three times as long as broad . . 17 

13 Edge of uncus lobe invaginated to form shallow pocket . . . . .14 

- Edge of uncus lobe not invaginated . . . . . . . .15 

14 Juxta approximately one-quarter length of valve (PI. 7, fig. 48) . amboinensis (p. 269) 

- Juxta more than half length of valve (PI. 7, fig. 46) . . . cinerascens (p. 267) 

15 Conspicuous inwardly directed elongate process from juxta or valve (PI. 7, fig. 49) 

errantia (p. 270) 

- Valve or juxta without inwardly directed process . . . . . . 16 

16 Tip of uncus lobe an inwardly curved spine (PL 8, fig. 52) . . albipuncta (p. 275) 

- Tip of uncus lobe broad, truncate (PI. 7, fig. 45) .... distracta (p. 269) 

17 Juxta extended dorsally into two very long, slender, curved processes, longer than 

the uncus ............. 18 

- Juxta not extended into elongate processes ....... 19 

18 Dorsal process from membrane between valve and juxta rectangular, with row of 

spines along distal edge ........ rigida (p. 277) 

- Dorsal process from membrane between valve and juxta broad, semi-ovate, inner 

surface covered with spines ....... baliomicta (p. 280) 

19 Uncus lobes with sharp tip, row of fine spines on internal surface (PI. 8, fig. 51) 

chalcites (p. 274) 

- Uncus lobes blunt, without fine spines on internal surface ..... 20 

20 Uncus approximately 1-3 times as long as wide; aedeagus 1-4 mm long, with large 

carina at two-thirds and two small apical thorn-like carinae (PI. 9, fig. 54) 

parallela (p. 281) 

- Uncus twice as long as wide; aedeagus 1 mm long, with small subterminal carina 

(PI. 9, fig. 57) ......... . insularia (p. 285) 



Key to female Tinissinae based on genitalia 
Females of Leptozancla and Tinissa bakeri, chaotica, errantia and ruwenzorica are unknown. 

[ African species ............ 

- Indo-Australasian species .......... 5 

1 Eighth tergite with transverse ridging, anterior margin with pouch cultellata (p. 296) 

- Eighth tergite without transverse ridging or anterior pouch. .... 3 
} Ostium at right angles to plane of eighth sternite ; ductus bursae with large ventral 

sclerotized patch (PI. 16, fig. 97) . . . . . . polystacta (p. 294) 

- Ostium oblique to or parallel to plane of eighth sternite, ductus bursae without 

sclerotization ............ 4 



REVISION OF THE TINISSINAE 265 

4 Ostium parallel to plane of eighth sternite, half as long as sternite; ductus bursae 

with fine rugose pimpling (PI. 16, fig. 98) .... poliophasma (p. 290) 

- Ostium oblique to plane of eighth sternite, almost as long as sternite; ductus bursae 

without pimpling (PI. 16, fig. 94) ...... spaniastra (p. 291) 

5 Eighth tergite with internal or external pouch or similar deep indentation, or 

sclerotized fold forming pouch at anterior margin ...... 20 

- Eighth tergite without pouch, any fold at anterior margin not sclerotized . . 6 

6 Ductus bursae with only about six regular constrictions (PI. 15, fig. 93) 

phrictodes (p. 289) 

- Ductus bursae with at least twenty regular constrictions ..... 7 

7 Distance from ostium to anterior tip of antrum one-half or less length of eighth 

sternite ............. 8 

- Distance from ostium to anterior tip of antrum three-quarters or more length of 

eighth sternite ............ 10 

8 Posterior margin of eighth tergite with deep U-shaped medial emargination . 9 

- Posterior margin of eighth tergite with only slight medial indentation (PI. 15, fig. 90) 

goliath (p. 287) 

9 Ventral lip of ostium with small, V-shaped medial emargination (PI. 14, fig. 87) 

insularia (p. 285) 

- Ventral lip of ostium with large, square emargination (PI. 15, fig. 91) 

kidukaroka (p. 287) 

10 Portion of antrum posterior to junction with ductus seminalis less than six times as 

long as wide ............ 12 

- Portion of antrum posterior to junction with ductus seminalis nine times or more as 

long as wide ............ 11 

11 Posterior projection of eighth sternite with dense dorsal patch of setae (PI. 16, fig. 96) 

torvella mysorensis (p. 294) 

- Posterior projection of eighth sternite quite smooth on dorsal surface (PI. 16, fig. 95) 

torvella torvella (p. 293) 

12 Conspicuous posterior lobe with short spines on seventh tergite (PI. 12, fig. 71) 

amboinensis (p. 269) 

- Seventh tergite without posterior lobe . . . . . . . . 13 

13 Anterior tip of antrum extending beyond tips of posterior apophyses (PI. n, fig. 67) 

krakatoa (p. 267) 

- Anterior tip of antrum not extending beyond tips of posterior apophyses . . 14 

14 Portion of antrum anterior to junction with ductus seminalis as long as or longer than 

posterior portion ........... 15 

- Portion of antrum anterior to junction with ductus seminalis shorter than posterior 

portion ............. 16 

15 Ventral surface of eighth tergite folded longitudinally to form accessory flaps either 

side of base of ovipositor (PI. 14, fig. 88) ..... dohertyi (p. 284) 

- Accessory flaps not present (PI. 11, fig. 68) .... cinerascens (p. 267) 

16 Posterior margin of eighth tergite with two deep lateral emarginations (PI. 11, fig. 69) 

distracta (p. 269) 

- Posterior margin of eighth tergite straight or with single medial emargination . 1 7 

17 Posterior margin of eighth tergite straight (PI. 14, fig. 86) . . indica (p. 282) 

- Posterior margin of eighth tergite with medial emargination . . . . 18 

18 Posterior margin of eighth sternite and ventral lip of ostium emarginated to about 

one half (PI. 14, fig. 89) ...... . transversella (p. 283) 

- Posterior margin of eighth sternite and ventral lip of ostium emarginated to less than 

one-quarter ............ 19 

19 Ostium oblique to surface of eighth sternite (PL 13, fig. 83) . . rigida (p. 277) 

- Ostium at right angles to surface of eighth sternite (PI. 13, figs 82, 84) 

baliomicta (p. 280) 



266 G. S. ROBINSON 

20 Conspicuous posterior lobe with short spines from seventh tergite . . . 21 

- Seventh tergite without posterior lobe . . . . . . . .22 

21 Antrum narrowed medially (PI. 12, fig. 76) .... araucariae (p. 273) 

- Antrum expanded medially (PI. 12, fig. 77) ..... chalcites (p. 274) 

22 Antrum straight-sided, extending beyond posterior third of anterior apophyses . 23 

- Antrum, if straight-sided, not extending beyond posterior third of anterior apophyses 24 

23 Pouch in eighth tergite more than three times as broad as antrum (PI. 12, fig. 79) 

palmodes (p. 272) 

- Pouch in eighth tergite approximately same width as antrum (PI. 12, fig. 75) 

eumetrota (p. 270) 

24 Eighth sternite heavily sclerotized and with deep emarginations either side of 

antrum and on anterior margin (PL 12, fig. 74) . . . convoluta (p. 272) 

- Eighth sternite with normal anterior margin . . . . . . .25 

25 Antrum straight, sometimes tapering, extending beyond anterior margin of eighth 

sternite ............. 26 

- Antrum, if straight, not extending beyond anterior margin of eighth sternite . 27 

26 Eighth sternite with deep medial, transverse fold, posterior half of sternite projecting 

beyond edge of fold (PI. 13, fig. 81) .... philippinensis (p. 277) 

- Eighth sternite not folded (PI. 13, fig. 80) ..... polysema (p. 276) 

27 Antrum bulbous, maximum diameter about half width of eighth sternite at widest 

point of antrum (PI. 12, fig. 78) ...... albipuncta (p. 275) 

- Antrum diameter one-third or less width of eighth sternite measured in same plane 

as antrum diameter ........... 28 

28 Pouch in eighth tergite one eighth from anterior margin, as broad as maximum 

width of antrum (PI. 15, fig. 92) ....... insignis (p. 288) 

- Pouch at anterior margin of eighth tergite, more than twice as broad as maximum 

width of antrum (PI. 14, fig. 85) ...... . parallela (p. 281) 



TINISSA Walker 

780. Type-species: 

Tinissa torvella Walker, 1864, ibidem 29 : 780, by monotypy. 
Polymnestra Meyrick, 1927, Exot. Microlepidopt. 3 : 331. Type-species: Polymnestra perilithias 

Meyrick, 1927, ibidem 3 : 331, by monotypy. [Synonymized by Gozmany & Vari, 1973 : 85.] 
Tinissa Walker; Meyrick, 1928 : 424 [redescription]. 
Tinissa Walker; Fletcher, 1929 : 182 [type-species]. 
Polymnestra Meyrick; Fletcher, 1929 : 224 [type-species]. 
Tinissa Walker; Diakonoff, 1955 : 127 [key to Papuan species]. 
Tinissa Walker; Diakonoff, 1967 : 278 [key to Philippine species]. 
Tinissa Walker; Gozmany & Vari, 1973 : 85 [redescription]. 

Mandible present; proboscis present, almost as long as second segment of labial palpus, 
absent in T. polystacta. Maxillary palpus usually three-segmented but with five segments 
in T. polystacta and T. cultellata. Antennae extending to about one-half of forewing costa, 
elongate cilia in male (PI. 1, fig. 1), shorter cilia in female (PI. 1, fig. 2), cilia very long in T. 
chaotica (<J) and reduced in length or absent in T. ruwenzorica, poliophasma and spaniastra. 
Fore wing (Text-fig. 3) somewhat rectangular, 5-15 mm in length; M present or absent in 
discal cell; R 5 to apex; i? 4 and i? 5 separate, stalked or completely fused. Elongate oval patch 
(PI 1, fig. 3) of small, flat, ovate scales (PI. 1, fig. 4) on ventral surface of fore wing between 
A x + 2 and posterior margin. Hind wing with patch of rough, pale scales opposite fore wing 
patch, anterior to Sc + R 1 . Venation usually complete but M sometimes absent from cell. 
Hind tibia with conspicuous distal hair tuft, first hind tarsal segment smooth-scaled (with 
rough, raised scaling in T. spaniastra). 



REVISION OF THE TINISSINAE 267 

Genitalia <$. Corema present in eighth segment of all but three species. Saccus large and 
triangular or long and slender. Uncus separated by membrane into pair of lobes attached 
to vinculum by narrow membranous band at anterior margin, fused to vinculum in T. cultellata 
and polystacta. Valve more or less reduced in most species; most species with digitate process 
arising from membrane between valve and juxta. Labides present in most species, a pair 
of (often fused) lobes dorsal to aedeagus, in several species extended to form ventral lobes. 

Genitalia $. See description for Tinissinae (p. 261) (the female of Leptozancla, the only 
other known genus of the subfamily, is unknown). 

Remarks. See remarks for Tinissinae (p. 263). 

Tinissa krakatoa sp. n. 

(PL 2, fig. 5; PI. ii, fig. 67) 

$ unknown. 

9- 18 mm. Head white. Labial palpus whitish, second segment brown above in basal 
half, tuft whitish with few brown streaks; terminal segment with brown subapical spot on 
dorsal surface, line of brown scales from apex to two-thirds on ventral surface. Thorax and 
tegula white, tegula brown anteriorly. Fore wing light ochreous brown heavily marked with 
whitish-ochreous coalescing transverse striae, termen darker; fringe light brown. Hind wing 
light grey-brown with a purplish sheen; fringe lighter, scales whitish at base. Legs whitish, 
hind tibial tuft ochreous cream. Fore leg light brown above, tibia and first two tarsal segments 
with deep brown streak on inner face. Mid tibia with two oblique light brown streaks on 
outer face, first tarsal segment with brown dot above. First hind tarsal segment with brown 
dot above. All spurs with brown dot below apex. 

Genitalia $ (PI. 11, fig. 67). Eighth sternite shallow with deep posterior medial emargina- 
tion to ostium. Ventral margin of ostium concave. Antrum funnel-shaped posteriorly, 
divided at three-quarters by membranous ring at point of junction with ductus seminalis. 
Anterior portion of antrum elongate, narrow, parallel-sided, almost reaching tips of apophyses 
anteriores. Eighth tergite tapered posteriorly, with small V-shaped medial emargination at 
tip. 

Remarks. T. krakatoa differs from T. cinerascens and T. amboinensis in the 
much longer anterior portion of the antrum which does not reach the tips of the 
apophyses posteriores in amboinensis or cinerascens. The female is smaller and 
paler than that of amboinensis. 

Distribution. Indonesia (Rakata I. [Krakatau]). 

Material examined. 

Holotype$, Krakatau: iv. 1933 (Dammerman) (genitalia slide no. 19750; BMNH). 

Tinissa cinerascens Meyrick 

(PI. 2, figs 6, 7; PI. 7, figs 46, 47; PI. 11, fig. 68) 

Tinissa cinerascens Meyrick, 1910, Trans, ent. Soc. Lond. 1910 : 476. LECTOTYPE 9> New 
Guinea: Louisiade Archipelago, Rossel I., 1905 (Meek) (genitalia slide No. 19786; BMNH), 
here designated [examined]. 

Tinissa cinerascens Meyrick; Diakonoff, 1955 : 128 [key]. 

<J. 13-15 mm. Pattern similar to $ but generally lighter, dark spots on fore wing costa 
not as frequent. Hind wing light greyish-ochreous, darker towards apex with some white 



268 G. S. ROBINSON 

maculae, the largest subapical on costa. Fore and hind wing fringes ochreous-whitish, tinged 
with grey at fore wing termen. 

9- 19-23 mm. Head and labial palpus whitish, vertex tufts light brown; second segment 
of labial palpus dark brown above, tuft mixed white and brown, third segment with few brown 
scales at base, oblique blackish brown band at one half. Antenna ochreous-whitish, first 
segment with brown scales above, scape white. Thorax and tegula whitish, tegula brown 
anteriorly and with broad brown medial band. Fore wing light ochreous brown heavily marked 
with ochreous-whitish such that brown ground colour remains as scattered transverse striae; 
terminal line dark brown. Fringe grey, scales at termen brownish basally. Hind wing brownish 
grey, paler maculae among darker spots at apex; fringe ochreous-whitish. Legs whitish, 
hind tibial tuft ochreous. Fore leg dark brown above to first tarsal segment, second and fourth 
segments with dark brown rings. Mid tibia with two oblique dark brown bands on outer 
face, first and fourth tarsal segments ringed with dark brown at distal end. Hind tarsal seg- 
ments one to four marked with dark brown above. All spurs with dark brown spot below 
apex. 

Genitalia ^ (PI. 7, figs 46, 47). Corema present in eighth segment. Saccus narrow, elon- 
gate, expanded anteriorly. Uncus broad, tips rounded with small lateral invagination. Sub- 
scaphium broad, setose posteriorly. Valves triangular, juxta forming conspicuous posterior 
lobe fused to each; process from membrane between valve and juxta present as lobe at tip of 
each valve. Aedeagus narrow, elongate, without carina. 

Genitalia O. (PI. 11, fig. 68). Eighth sternite rather square, rounded either side of terminal 
ostium. Ventral margin of ostium with deep U-shaped emargination. Antrum funnel- 
shaped posteriorly, divided at one half length of eighth sternite by narrow, membranous ring 
at point of junction with ductus seminalis; anterior portion of antrum narrow, straight-sided, 
2-5 times as long as posterior. Ductus bursae narrow, regular constrictions closely spaced. 
Eighth tergite truncate. 

Remarks. The invagination of the tip of the uncus and the presence of a process 
between the valve and juxta separate males of this species from those of T. distracta. 
Females may be separated by their more elongate posterior portion of the antrum, 
narrow ductus bursae and truncate eighth tergite. Females lack the lobe from 
the seventh tergite present in T. amboinensis , the male of which has a much smaller 
juxta than T. cinerascens. 

Distribution. Buru, New Guinea (Waigeo I., New Guinea, St Matthias Group, 
Louisiade Archipelago, Green Is. -Nissan I.), Australia (Queensland). 

Material examined. 

Paralectotypes, 2 $, 2 $, data as lectotype, 1 <$ as lectotype but Sudest [Tagula] I. 
(genitalia slide nos. 12288, 19723, 19787; BMNH). 

New Guinea: i £, Waigeo I., Camp Nok, 2500', iv. 1938 (Cheesman) (genitalia 
slide no. 19726); 1 $, Hydrographer Mts, 2500', iii. 1918 {Eichhorn); 1 $, Milne 
Bay {Meek) (genitalia slide no. 19725); 2 $, Louisiade Archipelago, St Aignan L, 
1897 {Meek) (genitalia slide no. 19711); 1 ?, St Matthias I., vi. 1923 {Eichhorn) 
(genitalia slide no. 19724); 1 <$, Green Is., Nissan I., viii-ix. 1924 {Eichhorn) (genitalia 
slide no. 19718). 

Australia: 4 $, Queensland, Cedar Bay, 1894 {Meek) (genitalia slide nos. 12287, 
19794). 

Buru: 2 $, below 1000', 1892 {Doherty) (genitalia slide nos. 12117, 19727). 
All specimens in BMNH. 



REVISION OF THE TINISSINAE 269 

Tinissa distracta Meyrick 
(PI. 2, figs 8, 9; PI. 7, fig. 45; PI. ii, fig. 69) 

Tinissa distracta Meyrick, 1916, Exot. Microlepidopt. 1 : 615. LECTOTYPE £, India: Assam, 
Khasi Hills, xii. 1906 (genitalia slide no. 19779; BMNH), here designated [examined]. 

£. 15-16 mm. Pattern similar to $ but fore wing even lighter, dark line at termen interrup- 
ted. Hind wing silky white, few dark brown maculae near apex, fringe white. 

§. 18-20 mm. Head and labial palpus whitish with few brown scales in vertex tufts; 
upper surface of second segment of labial palpus brown, brown subapical band on third segment. 
Antenna white, brown scales above on first segment. Thorax and tegula whitish, tegula 
brown anteriorly. Fore wing light ochreous brown almost obliterated by ochreous whitish 
fine transverse striae with larger spots at costa and end of cell, dark brown line at termen; 
fringe whitish, grey-brown at termen, terminal scales brown at bases. Hind wing light grey, 
whitish maculae towards apex; fringe whitish. Legs whitish, tuft of hind tibia tinted light 
ochre. Fore tarsi blackish brown above but third and fifth segments white. Mid tibia with 
two small blackish brown streaks on outer face; first and fourth tarsal segments banded with 
blackish brown distally. Hind tarsus with blackish brown bands at end of first and covering 
fourth segment, some brown at tip of third and proximal end of first segment. All spurs with 
blackish brown dot below apex. 

Genitalia^ (PI. 7, fig. 45). Corema present in eighth segment; saccus elongate, narrow, 
clubbed at anterior end, more than twice as long as uncus. Uncus broad, somewhat truncate. 
Valve large, fused with juxta which forms posterior and inner surface of fused complex. Juxta 
with two small conical projections on inner face, inner edge serrate below apex; labides present 
as pair of lightly sclerotized lobes against ventral surface of anellus. Aedeagus slender, elongate, 
two very small carinae below apex. 

Genitalia § (PI. 11, fig. 69). Eighth sternite rectangular, slightly produced posteriorly 
either side of terminal ostium. Antrum a little longer than eighth sternite, divided by mem- 
branous ring at point of junction with ductus seminalis; posterior section tubular, anterior 
section tapered. Regular constrictions of ductus bursae coarse. Eighth tergite with two 
deep lateral emarg.' r ations and a very small medial emargination posteriorly. 

Remarks. T. distracta is allied to T. cinerascens and externally very similar, 
but female cinerascens lack lateral emarginations of the eighth tergite; the portion 
of the antrum anterior to the junction with the ductus seminalis is three times the 
length of the posterior portion, not half as long as in distracta. Male distracta 
lack the lateral invaginations of the uncus to form a pair of pouches present in 
cinerascens. 

Distribution. India, Bali. 

Material examined. 

Paralectotypes, 6 $, data as lectotype, x, xi, xii. 1906, hi. 1907(3) (genitalia 
slide nos. 19731, 19791; BMNH). 

India: i $, Assam, Khasi Hills, x. 1895 (Doherty) (BMNH). 

Bali: i £, 1896 {Doherty) (genitalia slide no. 19741; BMNH). 

Tinissa amboinensis sp. n. 

(PL 2, figs 11, 12; PL 7, fig. 48; PL 11, fig. 70; PL 12, fig. 71) 

$. 15 mm. Head, labial palpus and antenna whitish, brownish scales above eye and in 
tuft of second segment of labial palpus. Labial palpus brownish above on outer face, terminal 



270 G. S. ROBINSON 

segment with ill-defined outer brown spot at two-thirds. Thorax and tegula whitish, tegula 
light brown anteriorly. Fore wing light greyish brown, darker at termen, densely flecked with 
yellowish white; fringe light greyish ochreous. Hind wing yellowish white, brownish on veins 
and at apex; fringe cream. Legs damaged; mid tibia with two oblique light brown bars on 
external face, first tarsal segment with light brown medial band, similar band at tip of third 
and base of fourth. First and second hind tarsal segments (remainder lost) with light brown 
bands towards tips. All spurs with dark brown spot below apex. 

$. 23 mm. Head and labial palpus whitish, badly rubbed. Second segment of labial 
palpus greyish brown above and on outer face, tuft streaked with greyish brown ; third segment 
distally brown from one-half, basal brown spot on outer face. Antenna ochreous, first and 
second segment brown above, scape whitish. Thorax and tegula rubbed. Fore wing greyish 
brown with lighter, ochreous, transverse striae, white scales at base, series of large cream 
spots along costa; fringe greyish brown. Hind wing greyish brown, paler maculae at apex; 
fringe light greyish brown, scales whitish basally. Legs damaged, ochreous-whitish, hind 
tibial tuft dark brown. Mid tibia dark brown above, first tarsal segment with two brown spots 
above. Hind tarsus with brown band on first segment; fifth, fourth and tip of third segment 
brown. 

Genitalia $ (PI. 7, fig. 48). Corema present in eighth segment. Saccus narrow, elongate, 
anteriorly expanded. Uncus broad, tips rounded with small lateral invagination. Sub- 
scaphium broad. Valve large, juxta forming conspicuous lobe at tip. Inner lobe from mem- 
brane between valve and juxta with inwardly directed thorn-like protuberance. Aedeagus 
elongate, narrow. 

Genitalia $ (PI. 11, fig. 70; PL 12, fig. 71). Seventh tergite produced posteriorly to form 
conspicuous lobe clothed with spine-like scales. Eighth sternite tapered, rounded at tip and 
medially emarginate to ostium. Ventral margin of ostium shallowly emarginate. Antrum 
elongate, 1-5 times as long as eighth sternite, divided by membranous ring at point of junction 
with ductus seminalis at two-thirds posteriorly. Eighth tergite with small, shallow V-shaped 
emargination at tip. 

Remarks. The lobe from the seventh tergite separates the female of this species 
from that of T. cinerascens which is much paler and narrower- winged. Males 
differ from those of cinerascens in having much smaller juxtal lobes at the tip of 
the valves and a more elongate aedeagus. 

Distribution. Moluccas (Ambon I.). 

Material examined. 

Holotype $, Ambon I. (Amboyna): 1892 (Doherty) (genitalia slide no. 19737; 
BMNH). 

Paratype <$, data as holotype (genitalia slide no. 19798; BMNH). 



Tinissa errantia sp. n. 

(PI. 2, fig. 10; PL 7, fig. 49) 

[Tinissa baliomicta Meyrick; Diakonoff, 1967 : 279, figs 423, 776, 778 (partim - 1 $ only). 

Misidentification.] 

^. 13 mm. Head, labial palpus and antenna white, first antennal segment with some 
brown scales above. Second segment of labial palpus with brown longitudinal line above, 
few brown scales in tuft; apical segment missing. Thorax and tegula whitish, light brown 
anteriorly. Fore wing yellowish white, few light brown transverse striae, dark brown marks 
at termen, two oblique brown basal lines to one-half of wing (inner line obscured in illustration) ; 



REVISION OF THE TINISSINAE 271 

fringe whitish, light greyish brown at termen. Hind wing white, veins yellowish, dark brown 
spots at apex; fringe white. Legs whitish, hind tibial tuft light yellowish brown. Fore tibia 
brown above, first tarsal segment with brown medial spot above; fourth tarsal segment brown, 
brown scaling extending on to third and fifth segments. Mid tibia with two light brown 
oblique bands on outer face; first tarsal segment with medial brown band, similar band on tip 
of third and base of fourth segment. Hind tarsus with brown medial band on first segment, 
brownish streak on under surface from band to tip. All spurs with brown dot below apex. 

$ unknown. 

Genitalia^ (PI. 7, fig. 49; Diakonoff, 1967 : fig. 423). Corema present in eighth segment. 
Saccus exceptionally narrow and elongate, three times as long as valve. Uncus lobes narrow, 
truncate. Details of fusion of valve and juxta found impossible to interpret but juxta probably 
forming conspicuous inward and dorsally directed processes; small triangular inwardly directed 
process may be homologous with lobe from membrane between valve and juxta observed in 
T. cinerascens and other species. Labides forming pair of ventral lobes. Aedeagus as narrow 
as saccus and twice as long. 

Remarks. T. errantia differs from T. distracta and its allies in possessing elongate 
processes from the tip of the valve-juxta complex. The uncus lobes are narrower 
and more truncated than in distracta and its allies, the known males of which differ 
from errantia in having the saccus expanded anteriorly. The holotype of errantia 
was described by Diakonoff (1967 : 279) as the 'neallotype' of T. baliomicta, a 
species not closely allied to distracta. The illustrations by Diakonoff (1967: figs 
776, 778) of the right hand side of the holotype are totally misleading as the right 
fore wing is bent, the distal half in deep shadow and appearing black. 

Distribution. Philippines (Luzon I.). 

Material examined. 

Holotype <$, Philippines: Luzon I., Mt Makiling {Baker) (genitalia slide no. 
5265 [Diakonoff]; USNM, Washington). 



Tinissa chaotica sp. n. 

(PI. 3, fig- 13; PL 8, fig. 50) 

C J. 16 mm. Head rubbed. Labial palpus yellowish, tuft of second segment streaked with 
light greyish brown, second segment brown above; terminal segment with brown dot on outer 
surface, below apex, some brown scales scattered towards base. Antenna ochreous-whitish, 
cilia three times width of flagellar segments. Thorax and tegula rubbed, whitish-ochreous. 
Fore wing light brown with densely packed ochreous-whitish transverse striae, darker brown 
at termen; fringe brownish white. Hind wing white, yellowish on veins, greyish brown at 
termen; fringe whitish. Legs whitish, hind tibial tuft tinted ochreous, slightly greyish beneath. 
Fore tibia dark brown above, first tarsal segment with brown dot above. Mid legs rubbed and 
damaged. Hind tarsus with pale yellowish brown diffuse mark at three-quarters on first 
segment, fourth segment with few yellowish scales. 

9 unknown. 

Genitalia <J (PI. 8, fig. 50). Corema absent from eighth segment. Saccus elongate, rectan- 
gular. Uncus tapered, setose, each lobe terminating in two heavily sclerotized spines. Valve 
short, rounded, with flap-like inwardly directed lobe from tip. Juxta produced posteriorly 
into two sclerotized points with U-shaped medial emargination a little broader than diameter 
of aedeagus, dorsally extended into two enormous processes with clubbed and spined tips 



272 G. S. ROBINSON 

which almost reach apex of uncus in undissected preparation. Labides forming pair of slender, 
elongate lobes dorsal to aedeagus, one-half length of uncus. Aedeagus slender, truncate. 

Remarks. Although externally similar to T. errantia, distracta and their allies, 
chaotica differs from all other known Tinissa species in having four tips to the 
uncus. The twin-pointed juxta and spined arms are similarly unusual as is the 
absence of coremata. This species may be distinguished externally by its very 
long antennal cilia. 

Distribution. Moluccas (Ambon I.). 

Material examined. 

Holotype <$, Ambon I. (Amboyna): 1892 (Doherty) (genitalia slide no. 12286; 
BMNH). 

Paratypes. 3 J, data as holotype (BMNH). 

Tinissa convoluta sp. n. 

(PI. 3, fig. 14; PI. 12, fig. 74) 

£ unknown. 

$. 20 mm. Head whitish. Labial palpus whitish, second segment brown above, tuft 
with few dark brown scales distally; apical third of terminal segment dark brown. Antenna 
ochreous white, scape white. Thorax and tegula very light brown, white scales on thorax 
posteriorly. Fore wing dark yellowish-ochreous with fine dark brown transverse striae, termen 
dark brown, some light marks on costa, notably towards apex; fringe dark greyish brown. 
Hind wing dark greyish brown, fringe lighter. Fore and mid legs white, light brown above; 
mid tibia with two oblique dull brown lines on outer face. Hind legs missing. 

Genitalia 9 (PI- I2 > n g- 74)- Eighth sternite broad, posteriorly emarginate to ostium, 
heavily sclerotized anteriorly, emarginate either side of antrum. Antrum lightly sclerotized, 
narrowed posteriorly, extended slightly beyond anterior margin of eighth sternite. Ductus 
bursae short. Eighth tergite shallowly emarginate posteriorly, posteriorly directed crescentic 
pouch with internal opening from near anterior margin; anterior margin produced into broad, 
shallow, T-shaped process covered by reflexed intersegmental membrane to form bilobed second 
pouch. 

Remarks. The anteriorly emarginate eighth sternite of convoluta is not observed 
in any other Tinissa species: the T-shaped pouch-forming process from the anterior 
margin of the eighth tergite is exclusively diagnostic. T. convoluta is externally 
similar to T. cinerascens but is slightly darker. 

Distribution. New Guinea (Bougainville I.). 

Material examined. 

Holotype $, New Guinea: Bougainville I. (Meek) (genitalia slide no. 19749; 
BMNH). 

Tinissa palmodes Meyrick 
(PL 3, fig. 15; PL 12, fig. 79) 

Tinissa palmodes Meyrick, 1917, Exot. Microlepidopt. 2 : 89. Holotype £, New Guinea: 

Setekwa R. (Snow Mts), 2-3000', 1916 (genitalia slide no. 19769; BMNH) [examined]. 
Tinissa palmodes Meyrick; Diakonoff, 1955 : 128 [key]. 



REVISION OF THE TINISSINAE 273 

g unknown. 

$. 29 mm. Head, thorax and tegula whitish; antenna light brown, scape whitish. Fore 
wing light ochre flecked with medium and dark brown scales, possibly to form transverse striae 
(but specimen very worn), large, lighter spots at costa, conspicuous spot at two-thirds, dark 
scales concentrated at end of cell, dark brown serrate terminal band; fringe scales whitish, 
grey-tipped, darker at termen. Hind wing very light grey-brown; fringe scales whitish tipped 
with grey-brown. Legs yellowish white; hind tibial tuft ochreous, dark brown at tips of 
distal scales. Fore femur and tibia brown above, tarsus with two brown dots. Mid tibia 
streaked with blackish brown on outer face, tarsus marked with light brown band at distal 
end of first and third segments. Hind tarsus tinted with light yellow-brown. All spurs with 
light brown band below apex. 

Genitalia $ (PI. 12, fig. 79). Eighth sternite rectangular, setose, deeply emarginate pos- 
teriorly; ostium slightly posterior to one half. Antrum long, funnel-shaped posteriorly, reach- 
ing tips of apophyses anteriores; ductus bursae short. Eighth tergite concave posteriorly, 
anterior margin folded into an extremely large, heavily sclerotized pouch with internal opening. 

Remarks. The excessively long antrum separates this from all other species 
with pouches in the eighth tergite, a feature which palmodes brings to the observed 
extreme. Allied to T. amboinensis which has no pouch but has a lobe from the 
seventh tergite. The holotype of palmodes has Cm x forked 1 mm from the termen 
in the right hind wing. 

Distribution. New Guinea. 

Material examined. 
Holotype only. 



Tinissa araucariae sp. n. 

(PL 12, figs 72, 76) 

[Tinissa chloroplocama Meyrick; Diakonoff, 1955 : 128. Misidentification.] 
<$ unknown. 

9- 23 mm. Head cream (badly rubbed). Labial palpus broken. Antenna whitish dor- 
sally, dull buff ventrally; scape white beneath. Thorax and tegula whitish, some light brown 
scales medially on thorax, anteriorly on tegula. Fore wing dark, dull brown, paler maculae 
discernible at costal margin, two small maculae towards apex, larger maculae at one-half, 
three-quarters and seven-eighths; blackish brown scales at apex running into fringe which is 
otherwise greyish brown. Hind wing and fringe light greyish brown, veins overlaid with darker 
brown scales. Legs badly damaged. 

Genitalia $ (PI. 12, figs 72, 76). Seventh tergite produced posteriorly to form conspicuous 
lobe bearing short, stout scales. Eighth sternite with shallow medial emargination: ostium 
terminal, ventral margin emarginate to one-fifth length of sternite. Antrum as long as eighth 
sternite, narrowed medially, sinuate anteriorly. Eighth tergite with concave posterior margin; 
posteriorly directed, heavily sclerotized pouch with ventral opening at one-quarter from anterior 
margin. Anterior margin of eighth tergite folded into single-lobed anteriorly directed pouch 
with dorsal opening. 

Remarks. T. araucariae is allied to T. chalcites but the ground colour of the 
fore and hind wing is much darker, the lobe of the seventh tergite shorter and 
broader, the eighth tergite posterior pouch not as deep, the ventral emargination 
of the ostium narrower than in chalcites; the antrum is not swollen medially as 



274 G. S. ROBINSON 

it is in chalcites. Although this specimen is worn and damaged it is quite distinct. 
Locality details are given by Diakonoff (1952 : 8). 

Distribution. New Guinea. 

Material examined. 

Holotype $, New Guinea: Araucaria Camp, 800 m, 26.iii.1939 (Toxopens) 
(genitalia slide no. L 03; RNH, Leiden). 

Tinissa chalcites sp. n. 

(PI. 3, fig. 16; PL 8, fig. 51; PI. 12, figs 73, 77) 

3. 23 mm. Pattern similar to $. 

9- 25-29 mm. Head light brownish yellow, tufts of darker scales near eyes. Labial 
palpus yellowish white; second segment partly light yellowish brown above, tuft white mixed 
with dark brown scales, terminal segment with light brown band at base and two-thirds. 
Antenna light ochreous brown, scape whitish. Thorax and tegula pale yellowish brown, 
thorax white posteriorly. Fore wing pale ochreous brown with medium brown transverse 
striae, denser markings along costa, termen and at one-quarter posteriorly; fringe medium 
brown with few lighter scales. Hind wing pale, dull yellowish brown, darker transverse striae 
at apex, dull brown at termen; fringe whitish, becoming grey towards apex. Legs yellowish 
white, hind tibial tuft light ochreous, distal scales tipped with greyish brown. Fore leg brownish 
above, two brown dots on upper surface of tarsus. Mid tibia with two oblique light brown 
bands on outer face. Two light brown dots on upper surface of hind tarsus. All spurs with 
brown band below apex. 

Genitalia $ (PI. 8, fig. 51). Corema present in eighth segment, eighth sternite markedly 
convex posteriorly. Saccus triangular, longer than wide. Uncus elongate, tips sht-rp, pointing 
outwards, line of fine spines pointing inwards from dorsal surface. Juxta enormous, forming 
pair of converging horn-like processes. Valve reduced, simple. Transtilla present; dorsal 
and ventral pairs of labides with pair of quadrangular ventrolateral labides adjacent to base 
of valve. Aedeagus bent at one-quarter, stout, several nodular projections on distal surface, 
largest just below apex. 

Genitalia £ (PI. 12, figs 73, 77). Seventh tergite produced posteriorly to form conspicuous 
lobe bearing short, stout scales. Eighth sternite with posterior medial emargination to one- 
fifth; ventral margin of ostium emarginate to level of posterior margin of eighth sternite; 
surface oi eighth sternite recessed towards ostium. Antrum as long as eighth sternite, slightly 
sinuate and narrowed at anterior end, swollen medially. Eighth tergite shallowly emarginate 
posteriorly; posteriorly directed, heavily sclerotized pouch with ventral opening at one-fifth 
from anterior margin which itself is rolled into a single-lobed, anteriorly directed pouch with 
dorsal opening; between these two pouches a pair of shallow lateral lobes. 

Remarks. T. chalcites is larger and paler than T. araucariae and has a brassy 
appearance; it is also allied to T. albipuncta (which is smaller and darker than both 
chalcites and araucariae), the female of which species has no spined lobe from the 
seventh tergite and the male of which does not have the tips of the juxta curved 
inwards. 

Distribution. New Guinea. 

Material examined. 

Holotype $, New Guinea: Mambare R., Biagi, 5000', i-iv. 1906 {Meek) (genitalia 
slide no. 19735; BMNH). 

Paratypes. 1 $, 1 9, data as holotype (genitalia slide no. 19730; BMNH). 



REVISION OF THE TINISSINAE 



275 



Tinissa albipuncta sp. n. 

(PI. 3. fig- 17; PI- 8. fig- 52; PL 12, fig. 78) 

£. 23 mm. Pattern similar to $ but thorax and tegula white, streaked anteriorly with 
brown. Markings of labial palpus and legs accentuated in comparison with §. 

5- 22 mm. Head whitish, vertex light brown. Labial palpus whitish, second segment 
light brown above and on outer surface of tuft; terminal segment with pair of light brown 
bands. Antenna dull brown, scape white. Thorax and tegula light yellowish brown flecked 
posteriorly with white. Fore wing dark, dull brown marked with conspicuous white dots at 
margins, large white dot at end of cell, transverse striae towards base; fringe brownish, lighter 
at tornus. Hind wing light ochreous brown, darker on veins and at apex, paler maculae at 
apex; fringe whitish, scales brown at base. Legs whitish, hind tibial tuft ochreous, darker 
distally. Fore and mid leg marked with brown above, two oblique brown bands on outer 
surface of mid tibia. All tarsi banded with brown (?- partly obscured by fungal hyphac), 
spurs banded below apex. 

Genitalia £ (PI. 8, fig. 52). Corema present in eighth segment. Saccus narrow, elongate. 
Uncus elongate, tapered, tips hooked inwards. Juxta enormous, fused to posteriorly extended 
vinculum, terminal lobes with inner line of heavily sclerotized dentate projections. Valve 
reduced, simple. Transtilla present; pair of stout, elongate labides dorsal to anellus [tips of 
these broken in illustrated preparation]. Aedeagus narrow, curved and elongate; small, short 
carina arising one-fifth from apex, small thorn-like projection below apex. 

Genitalia $ (PI. 12. fig. 78). Eighth sternite with broad posterior emargination. Dorsal 
margin of ostium slightly concave, ventral margin with V-shaped emargination to level of 
dorsal margin. Antrum elongate, globose, membranous in narrow band posterior to junction 
with ductus seminalis; portion of antrum anterior to junction with ductus seminalis L-shaped, 
narrow. Eighth tergite with small posterior emargination, heavily sclerotized pouch at anterior 
margin with two lateral, heavily sclerotized flaps. 

Remarks. T. albipuncta is smaller and darker than T. chalcites or T. araucariae, 
the wing pattern distinctive. The $ juxta is better developed than in chalcites 
which lacks a carina on the aedeagus. The $ lacks the lobe from the seventh 
tergite present in chalcites and araucariae. 

Distribution. New Guinea. 

Material examined. 

Holotype $, New Guinea: Mambare R., Biagi, 5000', i-iv. 1906 (Meek) (genitalia 
slide no. 10721; BMNH). 

Paratype <$, data as holotype (genitalia slide no. 19770; BMNH). 



Tinissa eumetrota Meyrick 
(PI. 3, fig- 18; PL 12, fig. 75) 

Tinissa eumetrota Meyrick, 1926, Exot. Microlepidopt. 3 : 319. LECTOTYPE $, New Ireland: 

xii. 1923-i. 1924 (Eichhorn) (genitalia slide no. 19785; BMNH), here designated [examined]. 
Tinissa eumetrota Meyrick; Diakonoff, 1955 : 128 [key]. 

£ unknown. 

9- 20-22 mm. Head brownish, whitish tufts near eyes. Labial palpus whitish, second 
segment dark brown above, tuft streaked with dark brown, terminal segment with dark brown 
subapical band, basally some dark scales on upper surface. Antenna dark brown, scape 
whitish. Thorax and tegula dark purplish brown with white transverse medial band. Fore 



276 G. S. ROBINSON 

wing dark purplish brown with paler transverse striae, whitish marks along costa, at end of 
cell, conspicuous transverse mark from posterior margin at nearly one-half; fringe greyish 
brown. Hind wing greyish brown with purplish sheen, paler maculae towards apex; fringe 
greyish brown. Legs ochreous white, tuft of hind tibia brown distally. Fore leg dark brown 
above, three dark brown bands on tarsus, basal band narrow, inconspicuous. Mid tibia with 
two dark brown oblique bands on outer face. First segment of hind tarsus with broad, dark 
brown band. All spurs with dark brown band below apex. 

Genitalia $ (PI. 12, fig. 75). Eighth sternite triangular, setose; ostium terminal; U-shaped 
sclerotized ridge from dorsal side of antrum at one-half. Antrum cylindrical, as long as eighth 
sternite, separated into two sections of similar diameter and length by membranous ring at 
point of junction with ductus seminalis. Eighth tergite posteriorly truncate, more heavily 
sclerotized anteriorly with small, narrow but deep anteriorly directed pouch with external 
opening. 

Remarks. Of species with pouches in the eighth tergite, only T. palmodes 
and T. eumetvota have an elongate antrum. In 'palmodes the antrum is completely 
sclerotized; in eumetrota there is a membranous ring at the point of junction with 
the ductus seminalis. The anterior tip of the antrum does not reach the tips of 
the apophyses anteriores as in palmodes. 

Distribution. New Ireland. 

Material examined. 

Paralectotype $, New Ireland: xii. 1923-i. 1924 (Eichhorn) (abdomen missing; 
BMNH). 



Tinissa polysema Zagulajev 
(PL 3, fig. 19; PL 13, fig. 80) 

Tinissa polysema Zagulajev, 1972, Trudy zool. Inst. Leningr. 52 : 348, figs 17a, 17b. Holotype 
$, Java: Baleq-Takengon, 29. ix. (Roepke) (genitalia slide no. 22; RNH, Leiden) [examined]. 

0* unknown. 

9- 24 mm. Head yellowish white, vertex greyish brown. Labial palpus yellowish white, 
tuft of second segment streaked with grey, upper surface spotted with brown; terminal segment 
black, white at base and apex. Antenna cream, some greyish ochre scales. Thorax and 
tegula greyish brown, paler posteriorly, medial transverse white band. Fore wing greyish brown, 
cream spots concentrated basally and at posterior margin, large cream spot at end of cell, deep 
greyish brown markings at apex and middle of termen ; fringe greyish brown flecked with white. 
Hind wing light greyish brown spotted with ochreous white towards apex and along costa, small 
brownish black mark at apex; fringe cream, scales tipped with grey. Legs whitish, tufts of 
hind tibia ochreous. Fore leg greyish brown above, tarsal segments banded with greyish 
brown. Mid tibia with two oblique greyish brown bands, only first tarsal segment banded. 
Hind tarsal segments banded with greyish brown. All spurs banded with dark grey below 
apex. 

Genitalia $ (PI. 13, fig. 80; Zagulajev, 1972 : figs 17a, 17b). Eighth sternite rectangular, 
setose; ostium slit-like, anterior; sternite deeply recessed to either side of ostium margin. 
Eighth tergite with posterior medial emargination, anteriorly with a crescentic, posteriorly 
directed pouch. Antrum two-thirds length of eighth sternite. 

Remarks. T. polysema is allied to T. philippinensis but the light forewing 
markings are bolder. The $ genitalia of philippinensis have the eighth tergite 



REVISION OF THE TINISSINAE 277 

only shallowly concave, the eighth sternite shorter and deeply folded dorsal to the 
ostium which is set in an emargination of the resulting transverse ridge. 

Distribution. Java. 

Material examined. 
Holotype only. 

Tinissa philippinensis sp. n. 

(PI. 3, fig- 20; PL 13, fig. 81) 
[Tinissa baliomicta Meyrick; Diakonoff, 1967 : 280 (partim - 2 § only). Misidentification.] 

<$ unknown. 

§. 22-23 mm. Head yellowish white, vertex with admixture of dark brown scales. Labial 
palpus cream, tufts of second segment streaked with medium brown, terminal segment with 
broad brownish black band below apex. Antenna cream, some brown scales at base of scape 
and on first segment. Thorax and tegula greyish brown with transverse medial white band, 
white at posterior margin. Fore wing medium brown, yellowish ochre spots forming pattern 
of broken transverse lines, larger spots at margins and at apex of cell, deep brown markings 
at apex and along termen; fringe light greyish brown with some whitish streaks. Hind wing 
light greyish brown, paler maculae towards apex, apex with three dark brown spots at margin; 
fringe light greyish brown with some lighter streaks. Legs yellowish white, hind tibial tuft 
light greyish ochre. Fore leg greyish brown above, fourth and fifth tarsal segments banded 
greyish brown at articulation. Mid tibia with two oblique greyish brown bands, first tarsal 
segment banded. Hind tarsus with very pale brown band on each segment. All spurs banded 
with dark brown below apex. Outer spurs of hind tibia with elongate dark-tipped scales. 

Genitalia O. (PI. 13, fig. 81). Eighth sternite tapered, setose, deeply infolded at two-thirds 
to form transverse ridge with deep medial emargination which anteriorly forms ostial margin. 
Posterior third of eighth sternite lightly sclerotized. Eighth tergite with shallow medial 
emargination posteriorly (depth greater in paratype), anteriorly with a crescentic posteriorly 
directed pouch. Antrum two-thirds length of eighth sternite. 

Remarks. The fore wing fringe of T. philippinensis is darker, the fore wing 
markings not as pale nor as bold as in T. polysema. The antrum is more heavily 
sclerotized and narrower, the eighth sternite not rectangular and not as heavily 
sclerotized posteriorly as in polysema. 

Distribution. Philippines (Luzon I.). 

Material examined. 

Holotype $, Philippines: Luzon, Mt Makiling {Baker) (genitalia slide no. W 05; 
USNM, Washington). 

Paratype $, data as holotype (genitalia slide no. W 08; USNM, Washington). 

Tinissa rigida Meyrick 
(PL 4, figs 2i, 22; PL 8, fig. 53; PI- 13. fig- 83; Text-fig. 4) 

Tinissa rigida Meyrick, 1910, Trans, ent. Soc. Lond. 1910 : 477. Holotype <$, Indonesia: 
Kai [Kei] Is., 1895 (K.) (genitalia slide no. 19782; BMNH) [examined]. 

Tinissa heterograpta Meyrick, 1928, Exot. Microlepidopt. 3 : 425. LECTOTYPE 9. New 
Britain: Talasea, hi. 1925 (Eichhorn) (genitalia slide no. 19780; BMNH), here designated 
[examined]. Syn. n. 



278 G. S. ROBINSON 

Tinissa chloroplocama Meyrick, 1938, Trans. R. ent. Soc. Lond. 87 : 526. LECTOTYPE <-J, 
New Guinea: Papua, Kokoda, 1200', v. 1933 (Cheesman) (genitalia slide no. 19790; BMNH), 
here designated [examined]. Syn. n. 

Tinissa rigida Meyrick, T. heterograpta Meyrick, T. chloroplocama Meyrick; Diakonoff, 1955 : 127 

[key]. " 

(J. 15-21 mm. Similarly patterned to $ but hind wing paler, a few darker maculae at apex. 
Flagellum of antenna whitish above. Brown markings on fore leg darker, brown bands on 
second and fourth tarsal segments, first with brown streak above, brown streak on outer surface 
of tibia. Very broad brown bands at one-third and two-thirds of hind tarsus. 

9- 2 1-25 mm. Head whitish, vertex with admixture of light brown. Labial palpus whitish, 
second segment brown above and on outer surface, brown extended to outer scales of tuft, 
terminal segment with broad brown ring at two-thirds, on several specimens a basal brown 
spot on outer surface. Antenna brownish, scape and pecten white. Thorax and tegula dark 
greyish brown anteriorly, whitish in posterior half. Fore wing dark brown to blackish brown 
with whitish transverse markings, two conspicuous oblique light basal lines; fringe greyish 
brown, scales whitish basally. Hind wing greyish brown with a violet sheen; fringes lighter. 
Legs ochreous-whitish, hind tibial tuft light greyish-ochreous. Fore leg greyish above but 
second and tip of first tarsal segment whitish. Mid tibia with two broad brown oblique bands 
on outer surface; brown dots above in middle of first and on fourth tarsal segment. Hind 
tarsus greyish brown above. All spurs with broad brownish streak below apex. 

Genitalia <§ (PI. 8, fig. 53). Corema present in eighth segment. Saccus rectangular. 
Uncus tapered, tips heavily sclerotized. Juxta large, heavily sclerotized, with broad, trebly 
emarginate pair of flange-like inwardly directed processes. Dorsal surface of juxta giving rise 
to pair of enormous horn-like processes. Valve reduced, small and stout, with dorsal digitate 
setose process at one-half; rectangular process from membrane between valve and juxta with 
broad band of inwardly directed spines at tip. Labides a pair of rectangular plates against 
dorsal surface of anellus and curved ventrad. Aedeagus short; short, blunt process just below 
apex. 

Genitalia $ (PI. 13, fig. 83; Text-fig. 4). Eighth sternite long, sharply tapered to form 
conical double-tipped projection which carries oblique ostium. Antrum broad, short, divided 
by membranous ring at point of junction with ductus seminalis, short anterior portion displaced 
to left. Eighth tergite with shallow V-shaped posterior medial emargination, folded longi- 
tudinally to form pair of internal lobes which slightly overlap eighth sternite. 

Remarks. The anterior portion of the antrum of T. rigida is larger than in 
T. baliomicta in which the plane of the ostium is at right angles to the plane of the 
abdomen, not oblique as in rigida. The lateral folds of the eighth tergite are not 
well-developed in baliomicta. Males of baliomicta from New Guinea have the 
dorsal process from the membrane between valve and juxta broad and thickly 
spined, not rectangular and only spined at the tip as in rigida. T. parallela is 
allied to rigida but the $ eighth tergite has an anterior pouch and the antrum is 
smaller. The $ of parallela lacks the horn-like processes from the juxta present 
in rigida and baliomicta. 

Distribution. Indonesia (Kai Is.), New Guinea, Karkar I., New Britain, 
New Hanover, St Matthias I., D'Entrecasteaux Is, Louisiade Archipelago, Australia 
(Queensland) . 

Material examined. 

New Guinea: 2 $, Kumusi R., low elev., v-ix. 1907 {Meek) (genitalia slide no. 
19777); 2 $>, 1 cS, Aroa R. (Meek) (genitalia slide no. 19716); 2 <$, Hydrographer 



REVISION OF THE TINISSINAE 



279 










Figs 4-10. Female genitalia (eighth sternite) of Tinissa spp. 4. T. rigida Meyrick [lecto- 
type of heterograpta], New Britain. 5. T. baliomicta Meyrick, holotype, Philippines. 
6. T. baliomicta Meyrick, New Guinea. 7. T. parallela sp. n., holotype, Sumatra. 8. 
T. dohertyi sp. n., holotype, Ambon I. 9. T. kidukaroka sp. n., holotype, Borneo. 10. 
T. insignis Zagulajev, holotype, Java. 



2 8o G. S. ROBINSON 

Mts, 2500', ii. 1918 (Eichhorn Bros.) (genitalia slide no. 10715); 1 £, Astrolabe Bay, 
1800', 1926 (Dodd); 2 (J, 2 $, Milne Bay (Meek) (genitalia slide no. 19713); 1 $, 
Lae, 20.ii.1968 (Coode). Karkar I.: 3 ?, 1 <$, ii. 1914 (Meek) (genitalia slide nos. 
19712, 19714). New Britain: i $, Talasea, i. 1925 (Eichhorn) [paralectotype 
of heterograpta] (abdomen missing). New Hanover: i $, ii. 1923 (Meek) (genitalia 
slide no. 19742). St Matthias I.: 1 <$, vii. 1923 (Eichhorn) (genitalia slide no. 
19717). D'Entrecasteaux Is.: 3 <$, 1 $, Fergusson I., 1-9. x., 7-20. xi., 21. xi- 
18. xii., 23-31.xii.1894 (Meek) (genitalia slide no. 19781). Louisiade Archipelago: 
1 $, St Aignan I., 1897 (Meek) (genitalia slide no. 19710). 

Australia: 1 ^ 2 ?, N. Queensland, Kuranda, n. x., 12. x., 20. xii. 1904 (Dodd) 
(genitalia slide nos. 19772, 19773). 

All specimens in BMNH. 

Tinissa baliomicta Meyrick 
(PI. 4. %. 23; PI. 13, figs 82, 84; Text-figs 5, 6) 

Tinissa baliomicta Meyrick, 1928, Exot. Microlepidopt. 3 : 424. Holotype $, Philippines: 

Luzon, Mt Makiling, 1927 (Baker) (genitalia slide no. 19789; BMNH) [examined], 
Tinissa baliomicta Meyrick; Diakonoff, 1967 : 279 (partim - 8 9_ only). 

<J [New Guinea only]. 21-23 mm. Similarly patterned to 9_ but fore wing with pale markings 
broader; dark, diffuse but conspicuous brown spot beyond whitish mark at end of cell. Hind 
wing paler than in $, very light greyish brown in one example, whitish in other. Fore tarsus 
brown above only on distal half of first and on fourth segment. Distinct black dot in upper 
surface of tuft at tip of hind tibia. 

9. 22-32 mm. Head whitish, brown scales near eyes and posteriorly. Labial palpus 
whitish, second segment brown above and on external face, tuft mixed with brown externally; 
base of terminal segment with few brown scales above, brown band at two-thirds. Antenna 
brownish, first segment very dark, scape white. Thorax and tegula whitish, brown anteriorly. 
Fore wing medium to dark brown with ochreous-whitish transverse striae, white concentrated 
at end of cell and in conspicuous oblique basal line, darker brown dot at base of costa and 
several small dark dots at apex; fringe greyish brown, paler basally, whitish dots continued 
into fringe. Hind wing greyish brown, paler maculae at apex; fringe mixed greyish and light 
ochreous. Legs ochreous-whitish, hind tibial tuft light greyish brown. Fore tibia and first 
and fourth tarsal segments brown above. Mid tibia with two broad oblique brownish bands 
on outer face, first and fourth tarsal segments with brown spot above. Upper surface of hind 
tarsus predominantly blackish brown, whitish only on base of first and tips of second and 
third segments. All spurs with large greyish brown streak below apex. 

Genitalia^ [New Guinea only]. Similar to genitalia of T. rigida but deep V-shaped medial 
emargination in ventral edge of juxta. Dorsal process from membrane between valve and 
juxta adjacent to dorsal pair of labides broad, semi-ovate and thickly clothed with fine spines 
on inner surface. 

Genitalia 9. (PI- 13, fig. 84; Text-fig. 5 [Philippines]; PL 13, fig. 82; Text-fig. 6 [New Guinea]). 
Eighth sternite triangular, deep, narrow medial emargination posteriorly. Ostium protuberant, 
at right angles to plane of eighth sternite. Antrum short, broad, sharply tapered anterior 
to membranous zone at point of junction with ductus seminalis. Eighth tergite with V-shaped 
medial emargination posteriorly. 

Remarks. T. baliomicta is closely allied to T. rigida but in the 9. genitalia the 
ostium is at right angles to the plane of the eighth sternite, not oblique: the $ 
has a broader and larger dorsal process between the valve and the juxta than does 



REVISION OF THE TINISSINAE 281 

rigida, spines not restricted to the margins of the process; males of rigida lack 
the deep V-shaped medial emargination present in the ventral margin of the juxta 
of baliomicta. Females of baliomicta from New Guinea differ slightly from topo- 
typical females in the more terminal position of the ostium and the medially more 
bulbous eighth sternite. In the three New Guinean preparations the eighth tergite 
has internal longitudinal folds forming lateral lobes similar to those of rigida. 
In specimens from Biagi these are much reduced but in a specimen from the Cyclops 
Mts are well developed. In New Guinea baliomicta and rigida appear to be geo- 
graphically separated, rigida only being recorded from below 750 m and baliomicta 
only from above 1050 m. The series of baliomicta described from the Philippines 
by Diakonoff (1967 : 279) contains four species, T. philippinensis , T. errantia 
and T. bakeri and only eight females referable to baliomicta. 

Distribution. Philippines (Luzon I.); New Guinea. 

Material examined. 

Philippines: 9 $, Luzon, Mt Makiling (Baker) (genitalia slide nos. W 04, 5266 
[Diakonoff]; USNM, Washington, one specimen in RNH, Leiden); 1 $, Luzon, 
Benguet, Palali, 2000', 20.xii.1912 (Wileman) (genitalia slide no. 19734; BMNH). 

New Guinea: 5 $, 1 3, Mambare R., Biagi, 5000', i-iv. 1906 (Meek) (genitalia 
slide nos. 19728, 19751, 19776; BMNH); 1 $, Cyclops Mts, Mt Lina, 3500-4500', 
iii. 1936 (Cheesman) (genitalia slide no. 19793; BMNH); 1 £, Buntibasa dist., Kratke 
Mts, 4000-5000', vi. 1932 (Mayer) (genitalia slide no. 19775; BMNH). 



Tinissa parallela sp. n. 

(PL 4, figs 24, 25; PI. 9, fig. 54; PI. 14, fig. 85; Text-fig. 7) 

$. 19-20 mm. Similarly patterned to § but paler; vestiture of head wholly yellowish 
white but brown external streaks on tuft of labial palpus, two external brown marks on terminal 
segment; few brown scales anterior to eye. Thorax and tegula white, tegula brown anteriorly. 
Fore wing pale markings heavier than in ^1 hind wing whitish, light brown on veins and at 
margin. Legs whitish, hind tibial tuft greyish brown distally. Fore leg dark brown above, 
tarsus banded with dark brown on first and fourth segments. Mid tibia with two oblique 
light brown streaks, mid tarsus with two brown dots on upper surface. Hind tarsal segments 
each with blackish brown spot on upper surface. All spurs with blackish brown subapical 
band. 

$. 22-23 nim. Frons yellowish white, vertex light brown. Labial palpus whitish, second 
segment brown above, tuft streaked with dark brown, terminal segment with two dark brown 
dots on outer surface. Antenna dull brown, lighter beneath, scape white. Thorax whitish, 
some light brown scales anteriorly, tegula light brown flecked with white. Fore wing dull 
brown with cream maculae, pronounced oblique double basal line, maculae concentrated in 
medial fascia and beyond cell, conspicuous dark mark beyond three-quarters; fringe whitish 
streaked with light brown. Hind wing light greyish brown, paler maculae towards apex; 
fringe whitish. Legs badly damaged. 

Genitalia^ (PL 9, fig. 54). Corema present in eighth segment. Saccus triangular, truncate 
anteriorly. Uncus tips blunt, heavily sclerotized, finely ridged at apex. Juxta very large, 
angular, without processes. Valve reduced, comparatively lightly sclerotized, with pair of 
dorsal processes with apices bearing long hairs. Pair of triangular labides dorsal to anellus. 



2$2 G. S. ROBINSON 

Aedeagus curved, tapering, triangular process at two-thirds, two small thorn-like processes 
at tip. 

Genitalia $ (PI. 14, fig. 85; Text-fig. 7). Eighth sternite tapered, truncate, a deep, narrow 
medial emargination to one-half; either side of emargination ventral ridges rising to meet 
ostial margin. Ostium transverse, protuberant. Eighth tergite medially emarginate pos- 
teriorly, margin invaginated anteriorly to form pouch. Antrum very short, less than half 
length of medial emargination of eighth sternite. 

Remarks. T. parallela differs from both T. rigida and T. baliomicta in lacking 
horn-like processes from the juxta in the $ and possessing a pouch in the eighth 
tergite in the $. It differs from T. indica in lacking spined processes from the 
subscaphium in the g and not having the antrum displaced anteriorly to the left 
in the $. 

Distribution. Singapore, Sumatra. 

Material examined. 

Holotype $, Sumatra: Padang Bovenlanden, Batang Proepoe, 1500' (genitalia 
slide no. 19792; BMNH). 

Paratypes. Sumatra: i 9> Dempo, 4000', viii. 1923 (Brooks) (genitalia slide 
no. 19736; BMNH); 1 $, Bonam Dolok, Sibolga, 500 m, 3.xi.i93i (Mohr) (genitalia 
slide no. L 04; RNH, Leiden). Singapore: i <$ (Baker) (genitalia slide no. W 06; 
USNM; Washington). 



Tinissa indica sp. n. 

(PL 4) fig. 26; PL 9) fig. 55; PL 14, fig. 86) 

$. 25-30 mm. Similarly patterned to $. 

§. 26-30 mm. Head whitish, few brown scales near eyes. Labial palpus whitish, second 
segment and tuft dark brown above and on outer surface, terminal segment with broad brown 
band at a little beyond one-half, brown spot at base. Antenna brown, scape and pecten 
whitish. Thorax and tegula predominantly brown, thorax white for posterior two-thirds, 
tegula white for posterior third with anterior white dot. Fore wing purplish brown with con- 
spicuous white dots along costa, in posterior third, at base and at end of cell, finely speckled 
with lighter brown; fringe brownish. Hind wing light greyish brown with a violet sheen, 
lighter speckles concentrated at apex; fringe lighter. Legs ochreous cream, hind tibial tuft 
light greyish brown. Fore tibia and tarsus blackish brown above, cream at articulations and 
on fifth tarsal segment. Mid tibia with two broad oblique brown bands on outer face, all 
tarsal segments except fifth with large blackish brown spot above, hind tarsi similar. All 
spurs with large blackish brown spot below apex. 

Genitalia g (PI. 9, fig. 55). Corema present in eighth segment. Saccus triangular. Uncus 
broad ; tips truncate, heavily sclerotized and setose. Subscaphium broad, invaginated to form 
pair of digitate processes with spined tips. Juxta large, forming pair of fist-shaped processes. 
Valve short, conical; elongate process from membrane between valve and juxta. Transtilla 
present, broad; pair of finely scobinate labides dorsal to anellus. Aedeagus with long carina 
from two-thirds. 

Genitalia £ (PI- 14, fig- 86). Eighth sternite ventrally protuberant, tapered, with shallow 
medial emargination in ventral margin of ostium. Antrum broad, ductus bursae entering on 
left. Posterior region of ductus bursae with very fine regular constrictions, ductus broadening 
and constrictions coarser anteriorly. Eighth tergite truncate, slightly concave posteriorly. 



REVISION OF THE TINISSINAE 283 

Remarks. T. indica is allied to T. transver sella but the $ lacks a deep posterior 
emargination of the eighth sternite or tergite; the wing pattern is much darker. 
T. indica is also allied to T. bakeri but the male has a truncate, not pointed uncus; 
the juxta is not reduced to a pair of cygnate lobes. 

Distribution. India (Assam), Sikkim, Bhutan, Taiwan (Formosa). 

Material examined. 

Holotype $, Sikkim: 1800', v. 1897 (Dudgeon) (genitalia slide no. 19766; BMNH). 

Paratypes. India: 3 $, 4 $, Assam, Khasi Hills, various dates 1894-1906 (Doherty) 
(genitalia slide nos. 19756, 19767, 19768); 2 <$, Assam, Margherita, 1888, 1889 
(Doherty) (genitalia slide no. 19759); 1 <$, Assam, Khasi Hills, Cherra Punji, 1894 
(Doncaster) ; 1 $, Assam, Silchar, Cachar, i2.viii.'7; 1 $, Assam, Jorhat dist., Tocklai, 
6-12.xi.1919 (Fletcher). Sikkim & Bhutan: i £ (Knyvett). Bhutan: i $, 28. vii. 
(Dudgeon) (genitalia slide no. 19756). All paratypes in BMNH. 

Material excluded from paratype series. Taiwan (Formosa): 2 $, Kanshirei, 
1000', 20. iv., 27. vii. 1908 (Wileman) (genitalia slide no. 19763; BMNH). 



Tinissa transver sella (Walker) comb. n. 
(PI. 4- ng. 27; PL 14, fig. 89) 

Gelechia transvey sella Walker, 1864, List Specimens lepid. Insects Colin Br. Mus. 29 : 641. 
Holotype $, Sarawak: Saunders colln (badly damaged - abdomen missing; BMNH) 
[examined] . 

$. unknown. 

5. 22-32 mm. [Description based on specimen from Sulawesi.] Head whitish mixed 
with brown. Labial palpus whitish, first and second segment blackish brown above and on 
outer surface of tuft, terminal segment with dark brown band at two-thirds. Antenna brown- 
ish, basal two segments of flagellum darker above, scape whitish. Thorax and tegula [damaged] 
whitish, dark brown anteriorly. Fore wing dull brown traversed by ochreous whitish striae; 
fringe greyish brown mixed with white, bases of scales pale. Hind wing light greyish with paler 
maculae at apex; fringe greyish brown, bases of scales whitish. Mid and hind legs missing: 
fore tibia and tarsus blackish brown above, whitish on second and fifth tarsal segments. 

Genitalia § (PL 14, fig. 89) [Specimen from Sulawesi]. Eighth sternite triangular with 
deep, narrow, medial emargination to one-half in dorsal and ventral surfaces of posterior projec- 
tion ; ostium very narrow. Antrum broad posteriorly, sharply narrowed anterior to membran- 
ous zone at point of junction with ductus seminalis; anterior portion of antrum narrow, sclero- 
tization extended into first nine or ten regular constrictions of ductus bursae. Eighth tergite 
with shallow posterior medial emargination, longitudinally folded to form two internal lateral 
flaps either side of ovipositor as in T. rigida. 

Remarks. The holotype of transversella is so damaged as to defy positive charac- 
terization of this taxon of which I take the female from Sulawesi described above 
as representative. The latter specimen is similar to the remnants of the type and 
is not incompatible with the original description. The Sumatran specimen differs 
slightly from the Sulawesi female in the lighter markings of the labial palpus and 
shorter anterior portion of the antrum: additionally it lacks the internal flaps from 
the eighth tergite but this feature is also variable in T. baliomicta. T. transversella 



284 G. S. ROBINSON 

differs from T. indica in having a much more extensive series of light markings 
on the fore wing and having a medially emarginate eighth tergite; the regular con- 
strictions of the ductus bursae are coarser than in indica. 

Distribution. Sumatra, Borneo (Sarawak), Sulawesi. 

Material examined. 

Sulawesi (S.W. Celebes): i $, Goa Malino, 3600', vi. 1938 (Kalis) (genitalia 
slide no. 19732; BMNH). 

Sumatra: i $, W. Sumatra, Lebong Tandai, 1920-23 (Brooks) (genitalia slide 
no. 19733; BMNH). 



Tinissa dohertyi sp. n. 

(PL 4, fig. 28; PI. 14, fig. 88; Text-fig. 8) 

^ unknown. 

§. 20 mm. Head rubbed bare; labial palpus whitish, second segment brownish above and 
on outer face, terminal segment damaged. Antenna broken but scape dull whitish. Thorax 
and tegula whitish posteriorly, brown anteriorly. Fore wing medium brown with creamy 
white transverse striae. Hind wing brownish grey, darker spots at apex; fringe of anal angle 
light brown with greyish tint. Legs whitish, damaged or missing, but one fore leg intact though 
rubbed: first and fourth tarsal segments banded with brown. Hind leg with greyish-ochreous 
tibial tuft, tip of tuft dark grey; tarsus brown above; spurs with dark brown streak below 
apex. 

Genitalia § (PI. 14, fig. 88; Text-fig. 8). Eighth sternite produced posteriorly, with deep 
ventral keel and posterior medial emargination to nearly one-half. Ostium deeply sunk 
within posterior lobes of eighth sternite. Antrum slightly tapering from ostium, bulbous 
at point of junction with ductus seminalis; anterior portion narrow, expanded slightly at tip. 
Eighth tergite with posterior medial emargination and internal longitudinal folds forming 
lightly sclerotized lobes either side of ovipositor. 

Remarks. T. dohertyi is closely allied to T. transversella but the posterior tips 
of the eighth sternite are more elongate; the eighth tergite has a deeper posterior 
medial emargination; the antrum is narrower, sclerotization not continued into 
the regular constrictions of the ductus bursae; the keel of the eighth sternite is not 
as deep in transversella as in dohertyi. 

Distribution. Moluccas (Ambon I.). 

Material examined. 

Holotype $, Ambon I. (Amboyna): 1892 (Doherty) (genitalia slide no. 19740; 
BMNH). 

Tinissa bakeri sp. n. 

(PI. 9) fig. 56) 

[Tinissa baliomicta Meyrick; Diakonoff, 1967 : 279 (partim - 1^ only). Misidentification.] 

3*. Head whitish, mixed with brown on vertex. Labial palpus whitish, brown above, tuft 
of second segment mixed with grey. Antenna white, ochreous below, a few brown scales 



REVISION OF THE TINISSINAE 285 

above on scape and first two segments. Thorax and tegula dull brown with broad transverse 
whitish medial band. Fore wings missing. Hind wing whitish, grey-brown on veins, fringe 
whitish. Legs [glued to polyporus] whitish; fore tibia blackish brown above, first and fourth 
tarsal segments banded with blackish brown. Mid tibia with two broad oblique light greyish 
brown bands, first mid tarsal segment banded with light greyish brown. Hind tibial tuft 
distally light greyish brown, first tarsal segment with light greyish brown transverse band ; 
three distal segments of hind tarsus missing. Spurs with greyish brown streak below apex. 

9 unknown. 

Genitalia $ (PI. 9, fig. 56). Corema present in eighth segment. Saccus short, very broad. 
Uncus lobes tapered, tips sharp, slightly curved. Subscaphium narrow posteriorly, broadened 
anteriorly and invaginated ventrad to form pair of heavily sclerotized, finely spined digitate 
processes. Juxta reduced, with deep medial emargination to form V-shaped process, tips of 
juxta arms broadened with inwardly directed spine. Valves divided into two lobes by deep 
medial notch: outer, ventral lobe rounded at tip; inner, dorsal lobe more elongate, tip produced 
into ventrally directed spine. Pair of lobate labides dorsal to anellus. Aedeagus stout, 
tapering apically, curved. 

Remarks. T. bakeri is closely allied to T. indica, the only other known species 
with a similarly modified subscaphium, but in indica the juxta is very stout, forming 
a pair of club-like processes curved over the valves. The process from the membrane 
between valve and juxta present in indica may be homologous with the dorsal lobe 
of the valve of bakeri: in the latter species there is a ventral membranous zone 
between the dorsal lobe of the valve and the remainder of the valve but on the 
ventral and inner face sclerotization is continuous. 

Distribution. Philippines (Luzon I.). 

Material examined. 

Holotype <$, Philippines: Luzon I., Mt Makiling {Baker) (genitalia slide no. 
Loi; RNH, Leiden). 



Tinissa insularia sp. n. 

(PI. 5, figs 29, 30; PI. 9, fig. 57; PI. 14, fig. 87) 

[Tinissa chloroplocama Meyrick, 1938, Trans. R. ent. Soc. Lond. 87 : 526 (partim - 1 ^ only). 

Misidentification.] 
[Tinissa lorvella Walker; Diakonoff, 1948, Treubia 19 : 218. Misidentification.] 
[Tinissa torvella Walker; Diakonoff, 1967 : 279, figs 428-432, 775. Misidentification.] 

<$. 17-21 mm. Similarly patterned to 0- but ground colour of legs much lighter so that 
dark markings are accentuated. 

§. 19-30 mm. Head cream, few brown scales near eyes. Labial palpus whitish, second 
segment brown on outer face and on tuft; terminal segment with blackish brown band at one- 
half. Antenna brownish, first two segments with darker scales above, scape whitish. Thorax 
and tegula dull brownish, lighter posteriorly. Fore wing dull brown with irregular, small, 
paler spots forming indistinct transverse striae, spots larger, whitish, at end of cell, on costa 
and at termen and forming a large, distinct spot on posterior margin at one-half; fringe greyish. 
Hind wing greyish-ochreous, apex with few darker dots; fringe whitish, scales greyish ochre 
from a little beyond bases to one-half. Legs light ochreous with greyish tint, hind tibial tuft 
greyish brown at apex. Fore tibia and tarsus dull grey above, few whitish scales at articula- 
tions. Mid tibia with two broad, ill-defined, greyish oblique bands on outer face, tarsal segments 
each with greyish spot above. First hind tarsal segment banded with greyish brown, second 



286 G. S. ROBINSON 

to fifth segments with greyish brown spot above. All spurs with greyish brown streak below 
apex. 

Genitalia £ (PL 9, fig. 57). Corema present in eighth segment, elongate, extending to 
anterior edge of seventh segment. Posterior margin of eighth segment more heavily sclerotized 
than usual in this genus, rounded, with closely packed scale-insertions. Saccus triangular, 
short. Uncus elongate, clubbed at tips, with transverse ridges across tips; elongate setae on 
dorsal surface. Juxta broad, enlarged, projecting dorsad posterior to valves which are reduced 
except for short, blunt dorsal process. Process from membrane between valve and juxta 
flattened, ovate. Labides fused to form conical projection dorsal to anellus; sclerotization 
extended short way anteriorly across membrane between valves, ventrally to form square, 
sclerotized plate on basal wall of anellus. Aedeagus short, narrow, with very small, short 
carina below tip. 

Genitalia 0. (PL 14, fig. 87). Eighth sternite projecting posteriorly with small V-shaped 
medial emargination forming ventral lip of terminal ostium. Antrum very short, tapering 
anteriorly; junction with ductus seminalis at one-half, point of junction sclerotized. Ductus 
bursae tapering posteriorly, regular constrictions very fine. Eighth tergite with broad, moder- 
ately deep U-shaped medial emargination. 

Remarks. T. insularia is a distinctive species; the fore wing markings are 
diagnostic, the white marks at the end of the cell smaller and less conspicuous than 
in T. goliath but the pale marks at one-half of the posterior margin of the fore wing 
are larger. The £ genitalia are close to those of T. parallela but the uncus is more 
elongate, more pronouncedly clubbed, the aedeagus not as elongate, the carina 
situated near the apex, not at two-thirds. The $ paralectotype of T. chloroplocama 
(below) has deformed genitalia with gynandromorph characteristics, the seventh 
tergite and sternite much deeper than is normal in females of this genus. 

Distribution. Malaya, Borneo, Sumatra, Java, Celebes (Kabia I.), Moluccas 
(Halmahera, Ambon, Buru), Philippines (Luzon, Mindanao, Palawan), New Guinea 
(Papua, Karkar I.), Solomon Is (Rendova I., Rennell I.). 

Material examined. 

Holotype $, Java: 192- (Kalshoven) (genitalia slide no. 19743; BMNH). 

Paratypes. Malaya: i $, Perak, Padang Rengas - low country, 1891 (Doherty) 
(BMNH); 1 <$, Penang, Waterfall v., 14. hi. -18. iv. 1898 (Curtis) (genitalia slide no. 
19774; BMNH). Borneo: i $, Sarawak, 1961-2 (Wallace) (BMNH); 1 $, Sarawak, 
1908 (W.B.) (genitalia slide no. 19747; BMNH); 1 9. Dutch West Borneo, 85 miles 
above Pontianak, Sanggan, iv. 1909 (Simons & Meligan) (BMNH). Java: 2 $, 
192- (Kalshoven) (BMNH). Celebes: i g, Kabia I. [Seleyer], Somarisi, 1660', 
xii. 1938 (Kalis) (genitalia slide no. 19739; BMNH). Moluccas: 2 $, Halmahera, 
1892 (Doherty) (genitalia slide nos. 19738, 19746; BMNH); 1 $, Ambon I., 1892 
(Doherty) (genitalia slide no. 19323; BMNH); 1 $, Buru, Station 9, 30.vi.1921 (Toxo- 
peus) (genitalia slide no. L 02; RNH, Leiden). Philippines: 3 $, 12 <J, Luzon I., 
Mt Makiling (Baker) (genitalia slide nos. Woi, W03, 5264, 5274; USNM, Washington 
[but 1 $ in RNH, Leiden and 1 $ in BMNH]; 1 $, Palawan I., Puerto Princesa, 
x. 1925 (genitalia slide no. 5263; USNM, Washington); 1 $, Mindanao I., Mt Apo 
School, 15 km SW. of Davao, 500 m, 22-31. x.1965 (Davis) (USNM, Washington). 
New Guinea: i $, Papua, Mafulu, 4000' (Cheesman) (genitalia slide no. 19788; 
BMNH) (paralectotype of chloroplocama Meyrick); 1 <$, Karkar I., ii. 1914 (Meek) 



REVISION OF THE TINISSINAE 287 

(BMNH). Solomon Is.: i <$, Rendova I. (Meek) (genitalia slide no. 19771; BMNH); 
1 $, Rennell I., Hutuna, 1-5. xi. 1953 (Bradley) (genitalia slide no. 19729; BMNH). 
Sumatra (North): 3 <$, 2 $, Dolok Merangier, 180 m, various dates 1969, 1970 
(Diehl) (MAK, Bonn); 1 $, Ketambe, 40 km NW. of Kutatjane, 300-500 m, 
10-18. vi. 1972 (Roesler & Kiippers) (MAK, Bonn). 



Tinissa goliath sp. n. 

(PL 5, fig. 31; PI. 15, fig. 90) 

$ unknown. 

9- 25 mm. Head white. Labial palpus whitish, brown on outer surface of second segment 
and tuft; terminal segment with brown dot on outer surface a little beyond one-half, few brown 
scales at base. Antenna brownish; scape and base of first segment white. Thorax and tegula 
light brown, tegula dark brown anteriorly. Fore wing greyish brown with a purplish sheen, 
marked with white spots, these concentrated at costa, posterior margin and end of cell and 
interspersed with smaller light greyish brown spots; fringe greyish brown. Hind wing light 
brownish grey, darker with scattered lighter maculae at apex; fringe light brownish grey, 
scales whitish at base. Legs whitish, hind tibial tuft greyish posteriorly. Fore leg heavily 
marked with dull brownish grey above, few whitish scales at articulations; mid leg similar, 
tibia with two grey oblique bands on outer face. Hind tarsus with two large dark greyish 
brown spots on first segment above, smaller spots on second, third and fourth segments. All 
spurs with greyish brown streak below apex. 

Genitalia 9 (PI- 15. fig- 90). Eighth sternite produced posteriorly with small but distinct 
posterior keel. Ostium terminal, very small; ventral margin with V-shaped indentation; 
dorsal margin deeply emarginate. Antrum extremely small, posteriorly bulbous, short. 
Ductus bursae broadened anteriorly, regular constrictions very fine and close-set. Eighth 
tergite protruding posteriorly; very small indistinct medial indentation in posterior margin. 

Remarks. The white fore wing markings of T. goliath are larger and more 
pronounced, especially the markings at the end of the cell, than in T. insularia 
in which the whitish mark at one-half of the posterior margin is much more pro- 
nounced. The ostium and antrum of goliath are much narrower than in insularia; 
the posterior margin of the eighth sternite is broader. The eighth tergite of goliath 
lacks the deep U-shaped posterior emargination present in insularia. 

Distribution. New Guinea. 

Material examined. 

Holotype $, New Guinea: [central W. Irian] Mt Goliath, 5-7000' (Meek) (geni- 
talia slide no. 19754; BMNH). 



Tinissa kidukaroka sp. n. 

(PI. 5, fig. 32; PI. 15, fig. 91; Text-fig. 9) 

cJ unknown. 

9. 21 mm. Head white, few brown scales near eyes. Labial palpus whitish, blackish 
brown above and on external face, second segment and tuft ; third segment missing. Antenna 
broken; scape white, first flagellar segment blackish brown above. Thorax whitish, tegula 
brown shading to cream posteriorly. Fore wing blackish brown patterned with white spots , 



288 G. S. ROBINSON 

spots particularly large at margins and coalesced at end of cell, basal two-thirds of wing badly- 
rubbed; fringe greyish brown, scales paler basally, apical and tornal white spots continued 
into fringe. Hind wing pale brownish grey, darker at distal margin, with a few pale apical 
spots; fringe greyish, scales paler basally. Legs whitish, hind tibial tuft light brownish grey. 
Fore leg damaged. Mid femur-tibia joint black above, tibia with two oblique greyish brown 
bands on outer face ; first tarsal segment with blackish medial band, third and fourth segments 
with blackish spot above. Hind tarsus similarly marked to mid tarsus. All spurs with blackish 
brown streak below apex. 

Genitalia § (PI. 15, fig. 91; Text-fig. 9). Eighth sternite with well developed ventral keel 
accommodating ostium and antrum; ventral lip of ostium squarely emarginate to nearly one- 
half length of posterior sclerotized portion of antrum ; dorsal margin of eighth sternite with 
shallow, V-shaped medial emargination. Antrum 8-shaped, anterior bulbous region mem- 
branous at point of junction with ductus seminalis; anteriorly a sclerotized ring posterior 
to point of junction with ductus bursae. Regular constrictions of ductus bursae very fine. 
Eighth tergite with broad U-shaped posterior medial emargination. 

Remarks. T. kidukaroka is closely allied to T. goliath but the white tornal 
markings of the fore wing are much heavier. The ostium is broader, its ventral 
lip much more deeply emarginate than in goliath) the antrum is longer and the 
eighth tergite has a deep medial emargination not present in goliath. 

Distribution. Borneo. 

Material examined. 

Holotype $, Borneo: N. Borneo, Kidukarok, 25.viii.1956 (Cambridge University 
Expedition) (genitalia slide no. 19744; BMNH). 



Tints sa insignis Zagulajev 
(PI. 5. ng. 33; PL 15, fig. 92; Text-fig. 10) 

Tinissa insignis Zagulajev, 1972, Trudy zool. Inst. Leningr. 52 : 350, figs 18a, 18b. Holotype 
§, Java: Nongkodjadjar, 6.viii.i934 (genitalia slide no. 23; RNH, Leiden) [examined]. 

$ unknown. 

9- 28 mm. Head whitish, vertex light brown. Labial palpus whitish, tuft of second 
segment dark, dull brown, few white scales distally; terminal segment denuded of scales. 
Antenna brownish, scape with few remaining ochreous cream scales. Thorax rubbed, ochreous 
cream, tegula dull brown, paler posteriorly. Fore wing dark purplish brown with ochreous- 
whitish maculae, most pattern detail obliterated; fringe mixed dull brown and brownish white. 
Legs brownish white. Fore leg blackish brown above, mid tibia with two blackish brown 
oblique stripes externally, fore and mid tarsal segments with darker bands, hind legs missing. 

Genitalia $ (PI. 15, fig. 92; Text-fig. 10). Eighth sternite tapered, setose, narrow posterior 
emargination with ridges either side rising to meet ventral margin of ostium at nearly one-half 
length of sternite: ostial margin almost perpendicular to and level with ridges. Antrum 
shorter than ridges with lobe-like internal projection from ventral wall. Eighth tergite with 
very deep medial emargination posteriorly, small, bilobed, anteriorly directed pouch at one- 
eighth anterior margin. 

Remarks. T. insignis is not closely aUied to any other Tinissa species but 
the $ genitalia resemble those of T. parallela. T. insignis is larger than parallela, 



REVISION OF THE TINISSINAE 289 

the pouch in the eighth tergite smaller, not at the anterior margin but at one-eighth 
from it. The ventral margin of the ostium is a sharp edge, not rounded as in 
parallela. 

Distribution. Java. 

Material examined. 
Holotype only. 

Tinissa phrictodes Meyrick 

(PL 5> figs 34, 35; PL 9, fig. 58; PL 15, fig. 93) 

Tinissa phrictodes Meyrick, 1910, Trans, ent. Soc. Lond. 1910 : 477. LECTOTYPE 9. Solomon 
Is.: Choiseul I., 1905 (Meek) (genitalia slide no. 19783; BMNH), here designated [examined]. 

£. 16-18 mm. Similarly patterned to $ but antenna whitish, first and second flagellar 
segment with few brown scales above. Fore wing with white markings much more numerous 
and close-set than in $; dark termen conspicuous. Hind wing greyish white, veins darker, 
apex greyish brown; fringe paler than in 9- Legs whiter than those of 9 so dark markings 
more conspicuous, hind tarsus with dark brown spots above on first to fourth segments, dark 
scaling extended to form band on first segment. 

9- 21 mm. Head whitish, few brown scales near eyes. Labial palpus whitish, second 
segment brown above and on outer face; outer surface of tuft brown; terminal segment with 
brownish streak from one-half on outer face, few brown scales at base. Antenna brownish, 
scape and first flagellar segment white. Thorax and tegula light brown, tegula with medial 
transverse white band. Fore wing light creamy brown, rather darker at termen, patterned 
with whitish spots, spots large at costa and posterior margin, coalescing at end of cell; fringe 
light grey, scales paler at base. Hind wing light brownish grey, apex darker, few pale subapical 
spots; fringe light grey, scales paler at base. Legs ochreous-whitish, hind tibial tuft brownish 
grey. Fore leg dark grey above, two triangular white marks on outer face of tibia, second and 
fifth tarsal segments white, some whitish scales on adjoining segments. Mid tibia with two 
diffuse, oblique greyish bands on outer face, brownish grey bands on first, third and fourth 
tarsal segments. Hind tarsus with first segment brownish from one third, few brownish 
scales at base. All spurs with brownish streak from below apex almost to base. 

Genitalia <$ (PI. 9, fig. 58). Corema absent from eighth segment. Saccus elongate, triangu- 
lar. Uncus lobes divergent, supported on broadened section of vinculum two-thirds as deep 
as saccus. Tips of uncus lobes turned inward, bearing line of dense, short spines. Subscaphium 
distinctly V-shaped. Juxta a narrow, heavily sclerotized ring ventral to anellus, nearly com- 
plete dorsally, forming pair of horn-like dorsal processes. Transtilla present, forming broad 
bar between valves dorsal to juxta processes. Valve elongate, tip heavily sclerotized, pointed, 
directed ventrad. Aedeagus short and stout. 

Genitalia 9 (PI- J 5. ng. 93). Eighth sternite almost square, posterior third setose; small, 
shallow V-shaped posterior emargination; ostium situated at three-quarters. Ventral lip of 
ostium with very deep, narrow, V-shaped medial emargination. Antrum extremely short. 
Ductus bursae broad, only five or six ill-defined regular constrictions posteriorly. Eighth 
tergite with broad, shallow V-shaped medial emargination posteriorly. 

Remarks. The <$ of phrictodes differs from other Tinissa species in the degree 
of dorsal broadening of the vinculum, possessing a V-shaped subscaphium and in 
the characteristic shape of the juxta and uncus. The 9 may be recognized by the 
long and tapering bursa copulatrix and the extremely short and inconspicuous 
section of the ductus bursae which is regularly constricted. The position of this 
species is obscure; the structure of the uncus is similar to that of T. polystacta 
but phrictodes has a three-segmented maxillary palpus whereas the palpus of poly- 



2go G. S. ROBINSON 

stacta is five-segmented. The $ specimen from New Ireland has a small sclerotized 
area on the wall of the bursa copulatrix. This is the only case of sclerotization of 
the bursa noted in this subfamily: it is considered to be a freak. 

Distribution. New Ireland, Solomon Is. (Choiseul, Rendova, Vella Lavella 
Is.). 

Material examined. 

Solomon Is.: i <$, Choiseul I., 1905 {Meek) (genitalia slide no. 19784; BMNH) 
[paralectotype] ; 1 <$, Rendova I. (Meek) (genitalia slide no. 19720; BMNH); 1 $, 
Vella Lavella I., ii-iii. 1908 (genitalia slide no. 19722; BMNH). New Ireland: 
1 $, xii. 1923 (Eichhorn) (genitalia slide no. 19745; BMNH). 

Tinissa poliophasma Bradley 
(PI. 5, ng. 36; PL 16, fig. 98) 

Tinissa poliophasma Bradley, 1965, Ruwenzovi Exped. ig?j2 2 : 115, figs 56, 204, 205. Holotype 
$, Uganda: Ruwenzori Range, Semliki Forest, 2850', 22.viii.-3.ix.1952 (Fletcher) (genitalia 
slide no. 8483 [Bradley]; BMNH) [examined]. 

Tinissa poliophasma Bradley; Gozmany & Vari, 1973 : 86, fig. 226 [redescription] . 

$ unknown. 

§. 19 mm. Head white with few brown scales near eyes. Labial palpus white, first and 
second segments brown above, tuft mixed with brown; terminal segment with brown spot on 
outer face a little beyond one-half. Antenna light ochreous, scape white, scales of pecten 
tipped with brown, first flagellar segment brown above; cilia extremely short. Thorax and 
tegula white, light brown anteriorly. Fore wing light brown, darker at termen, marked with 
close-set lighter transverse striae, conspicuous white marks at costa, conspicuous white basal 
fascia, large pale spot on posterior margin at one half; fringe ochreous, greyish brown at termen. 
Hind wing light golden brown with a greyish tint ; fringe whitish, scales tinted with light brown 
from little beyond base to more than one-half of their length. Legs (damaged) ochreous- 
whitish. Fore leg greyish-ochreous above to tip of tibia, greyish-ochreous spot on upper surface 
of first tarsal segment. Mid femur-tibia joint brownish, tibia with two light brown oblique 
bands on outer face, first and fourth tarsal segments with light brown spot above. Hind legs 
missing. 

Genitalia $ (PI. 16, fig. 98). Eighth sternite almost square; ostium from one half to tip 
of sternite, in same plane as surface of abdomen; ostium lip slightly protuberant. Antrum 
extremely short, about half length of ostium. Ductus bursae with regular constrictions very 
fine, rugose, rugosity appearing as overlapping plates under high ( x 250) magnification. Eighth 
tergite rounded posteriorly. 

Remarks. The wing pattern of this species differentiates it from all other 
known Tinissa species which lack the strongly defined white basal fascia of the 
fore wing. The extremely short antennal cilia of poliophasma are possibly evidence 
for alliance with T. rnwenzorica but the female of the latter species is unknown. 
This is the only Tinissa species in which the ostium is parallel with the abdomen 
surface; in all other known species it is at least oblique. 

Distribution. Uganda (Ruwenzori Range). 

Material examined. 
Holotype only. 



REVISION OF THE TINISSINAE 291 

Tinissa spaniastra Meyrick 
(PL 6, fig. 37; PI. 9, fig. 59; PI. 16, fig. 94) 

Tinissa spaniastra Meyrick, 1932, Trans, ent. Soc. Lond. 80 : 118. Holotype $, Ethiopia: 
Djem-Djem Forest, c. 8000', 9.x. 1926 (Scott) (genitalia slide no. 10176 [Gozmany]; BMNH) 
[examined]. 

Tinissa spaniastra Meyrick; Gozmany, 1969 : 290 [description of $ genitalia]. 

Tinissa spaniastra Meyrick; Gozmany & Vari, 1973 : 85, figs 223, 224 [redescription] . 

(J. 25 mm. Similarly patterned to $ but legs with all tarsal segments except first (as §) 
brown to at least two-thirds except for fifth tarsal segment which is white. Antennal cilia 
longer than in O. but still only one-half diameter of shaft. 

9- 20, 27 mm. Head cream, few brown scales near eyes. Labial palpus cream, second 
segment and tuft brown on outer surface; terminal segment ringed with dark brown at one-half. 
Antenna, including scape, brownish; first flagellar segment nearly black above; cilia very short. 
Thorax and tegula dark brown, thorax with few white scales posteriorly. Fore wing dark 
brown, darker at costa and termen, boldly marked with white spots, most conspicuous spot on 
costa at two-thirds; fringe brown. Hind wing light brownish ochre with numerous paler 
maculae, ground colour darker towards apex; fringe light ochreous. Legs ochreous to white, 
hind tibial tuft greyish brown. Fore leg dark brown above, some white scales at articulations, 
fifth tarsal segment white. Mid tibia with two broad, oblique brown bands on outer face, 
first tarsal segment with broad brown medial ring; remaining segments missing. First hind 
tarsal segment with rough brown scaling above for two-thirds; second and third segments 
brown basally, remainder missing. All spurs with dark brown streak below apex. 

Genitalia g (PI. 9, fig. 59). Corema present in eighth segment. Saccus triangular, elongate. 
Uncus lobes broad, tips produced into inwardly directed spines. Juxta forming pair of broad 
lateral projections with sharp dorsal tips; valve reduced, conical; process from membrane 
between valve and juxta long, finger-like, reaching tip of juxta. Labides fused and forming 
large hood-like projection dorsal to valves. Aedeagus slender, tapering, with line of fine 
spicules running full length, scattered at base and dense at tip. 

Genitalia $ (PI. 16, fig. 94). Eighth sternite long and narrow, line of fine spines at pos- 
terior margin; ostium almost as long as sternite, tapering posteriorly, angled obliquely to plane 
of sternite, ventral lip V-shaped. Antrum short, three-quarters length of eighth sternite, 
slightly tapering anteriorly, only lightly sclerotized at point of junction with ductus seminalis. 
Regular constrictions of ductus bursae very fine, minutely rugose. Eighth tergite rounded 
posteriorly with about twelve coarse spines at margin. 

Remarks. T. spaniastra is allied to T. poliophasma but the ostium, antrum 
and ductus bursae are broader; the ostium is not flush with the surface of the eighth 
sternite; the rugosity of the ductus bursae is much coarser in poliophasma. The 
ostium of spaniastra tapers posteriorly but is almost rectangular in poliophasma. 
The fore wing pattern of this species is sufficiently distinctive to be diagnostic, 
the major distinguishing feature being the large white costal spot. Gozmany's 
dissection of the holotype of this species is not really suitable for critical examination. 
The genital armature has been so badly crushed that I am loth to attempt to remount 
the preparation. Gozmany & Vari's (1973) description of the male genitalia of 
spaniastra should be ignored as all components within the vinculum with the excep- 
tion of the aedeagus are misidentified (see 'Morphology'). The 'cornuti' of the 
same authors are fine spicules on the outer surface of the aedeagus. 

Distribution. Ethiopia, Tanzania. 



292 g. s. robinson 

Material examined. 

Ethiopia: 2 $, Maraco, 20. ix., 30.x. 1915 (Kovacs) (genitalia slide no. 19795; 
BMNH). 

Tinissa ruwenzorica Gozmany 
(PL 6, fig. 38; PL 10; figs 60, 61) 

Tinissa ruwenzorica Gozmany, 1966, Acta zool. hung. 12 : 68, fig. 18. Holotype <§, Uganda: 
Ruwenzori Range, Mahoma River, 6700', 13-16. viii. 1952 {Fletcher) (genitalia slide no. 12013 
[Gozmany - remounted Robinson]; BMNH) [examined]. 

Tinissa ruwenzorica Gozmany; Gozmany & Vari, 1973 : 85, fig. 225 [redescription] . 

(J. 17 mm. Head whitish, few brown scales laterally. Labial palpus whitish, first and 
second segment and tuft flecked with dark brown on outer face; terminal segment with dark 
brown spot on outer face a little beyond one half. Antenna ochreous-whitish, first flagellar 
segment dark brown above; scape white; cilia extremely short. Thorax and tegula white, 
dark brown anteriorly. Fore wing dark blackish brown heavily striated with cream mixed with 
light brown, striations concentrated basally and in postmedial fascia, blackish terminal line, 
tornus almost uniformly blackish brown; fringe greyish, whitish at termen and mixed with 
brown posteriorly. Hind wing whitish, few dark maculae at termen; fringe whitish. Legs 
whitish, hind tibial tuft greyish brown. Fore leg dark grey above. Mid femur-tibia joint 
grey, tibia with two dark grey oblique bands on outer face, first tarsal segment with grey 
spots basally and at one-half above, tarsus otherwise grey above. Hind tarsus similarly 
patterned, dark grey raised scales basally on upper surface of first segment. All spurs with 
subapical brown ring. 

$ unknown. 

Genitalia <$ (PL 10, figs 60, 61). Corema present in eighth segment, corematal scales coarse, 
needle-like. Eighth sternite moderately sclerotized, produced posteriorly at corner to form 
slightly curved and tapering process. Saccus elongate, triangular. Uncus lobes short, rounded, 
each with digitate lateral projection. Juxta produced posteriorly, narrow, terminating in 
pair of clubbed, cygnate processes. Valve short, conical. Process from membrane between 
valve and juxta broad, flat, parallelogram-shaped, as long as valve. Labides fused, forming 
short projection dorsal to anellus. Aedeagus slender and tapering, small carina just below 
apex. 

Remarks. Although allied to T. torvella, possessing a similarly modified eighth 
sternite, ruwenzorica differs in having very short antennal cilia and in the modifica- 
tion of the juxta which is reminiscent of that of T. bakeri. The form of the uncus 
of ruwenzorica is distinct - as is the external appearance - from that of all other 
known Tinissa species. The original and subsequent descriptions of this species's 
genitalia are inaccurate, as are the illustrations. 

Distribution. Uganda (Ruwenzori Range). 

Material examined. 
Holotype only. 

Tinissa torvella Walker 

(PL 6, fig. 39; PL 10, figs 62, 63; PL 16, figs 95, 96) 

Tinissa torvella Walker, 1864 : 780. 

O*- 16-24 mm. Similarly patterned to $> but elongate scales of vertex all whitish, transverse 
whitish band across thorax and tegula broader. 



REVISION OF THE TINISSINAE 



293 



§. 20-32 mm. Head whitish, mixed with blackish brown near eyes and on vertex. Labial 
palpus whitish, second segment and tuft blackish brown on outer face, terminal segment with 
blackish brown spot on outer surface at base, blackish brown band at two-thirds. Antenna 
brownish, blackish above, scape white, scales of pecten tipped with black, first two flagellar 
segments black above. Thorax and tegula blackish brown with medial white transverse 
band. Fore wing blackish brown marked with numerous small white spots, spots larger and 
conspicuous at margins, in an ill-defined basal line and at end of cell; fringe blackish brown, 
scales paler basally, marginal white spots extended into fringe. Hind wing light brownish 
grey, whitish maculae at apex; fringe light grey mixed with off-white, pale apical spots extended 
into fringe. Legs greyish white, hind tibial tuft grey. Fore femur dark grey above, tibia 
and first tarsal segment grey; second and fourth tarsal segments banded with blackish. Mid 
tibia dark brownish grey above for proximal third; two oblique dark brownish grey bands on 
outer face; first tarsal segment with blackish medial band, fourth segment blackish. First 
and fourth hind tarsal segments banded with blackish. All spurs with long blackish streak 
below apex. 

Genitalia <$ (nominate subspecies only - PI. 10, figs 62, 63). Corema present in eighth 
segment. Eighth sternite produced posteriorly at corner to form conspicuous, finely spined, 
club-shaped projection. Saccus broad, elongate. Uncus arms narrow, tapering. Sub- 
scaphium broad anteriorly. Juxta dish-shaped, almost surrounding aedeagus, not closely 
appressed to valve. Process from membrane between valve and juxta elongate and conspicu- 
ous, internal surface setose. Valve tooth-shaped, reaching three-quarters length of process. 
Base of anellus sclerotized dorsally, possibly representing labis. 

Genitalia § (PI. 16, figs 95, 96). Eighth sternite broad anteriorly, abruptly narrowed at 
one-third or one-half and produced posteriorly as elongate, dome-like structure; dorsal durface 
of this 'dome' with or without setae, reflexed anteriorly to form shallow pouch, base of which is 
level with anterior tip of antrum or at one-half. Ostium terminal, small; antrum narrow- 
posteriorly, bulbous and only thinly sclerotized at point of junction with ductus seminalis. 
Anterior portion of antrum one-quarter length of posterior and three times as broad. Eighth 
tergite with deep U-shaped posterior medial emargination. 

Remarks. The modification of the male eighth sternite differentiates this 
from all other known Tinissa species with the exception of T. ruwenzorica in which 
the uncus lobes are rounded with a small but conspicuous lateral projection. The 
female genitalia are characterized by the large, dome-like posterior projection. 
Described material is fresh; older material is faded to a dull brown. This species 
is represented by two subspecies, described below. 

Distribution. Sri Lanka (Ceylon), India (Mysore). 



Tinissa torvella torvella Walker 
(PI. 6, fig. 39; PI. 10, figs 62, 63; PL 16, fig. 95) 

Tinissa torvella Walker, 1864, List Specimens lepid. Insects Colin Br. Mus. 29 : 780. Holotype 
,3\ [Sri Lanka (Ceylon):] 'Assam?' [erroneous] (genitalia slide no. 10344 [Gozmany] ; BMNH) 
[examined]. 

Genitalia $ (PL 10, figs 62, 63). See description above. 

Genitalia § (PI. 16, fig. 95). Differs from torvella mysorensis in that the eighth sternite 
is narrowed at one-third rather than one-half, dorsal surface of posterior 'dome' without setae, 
reflexed anteriorly to form shallow pouch, base of which is level with anterior tip of antrum, 
not at one half of antrum as in torvella mysorensis. Antrum shorter and narrower than in 
torvella mysorensis. 



294 G - s - ROBINSON 

Remarks. The nominate subspecies of T. torvella is differentiated from T. 
torvella mysorensis on characteristics of the $ genitalia as described above; it is 
not separable from torvella mysorensis by external characteristics. 

Distribution. Sri Lanka (Ceylon). 

Material examined. 

Sri Lanka [Ceylon]: i $, Maskeliya, viii. 1905 (de Mowbray) (USNM, Washington); 
1 $, Maskeliya, vi. 1905 (de Mowbray) (genitalia slide no. 19752; BMNH); 5 $, 1 <$, 
Kandy, 1600', iii, iv, v, viii, 189-, 1909, 1911 (Pole, Mackwood) (genitalia slide 
nos. 19753, 19758, 19799 [ wm gs]: BMNH); 1 $>, Peradeniya, viii. 1905 (Green) 
(BMNH); 2 $, W. Matale, 400', 189- (Mackwood) (BMNH); 1 $, 3 <J, Nawalapitiya, 
2000', 189- (Pole) (genitalia slide no. 19757; BMNH); 1 <$, Colombo, 1891 (Mackwood) 
(genitalia slide no. 19322; BMNH) ; 1 $, N. Chiga, 6000', v. 1902 (Mackwood) (BMNH) ; 
1 $, Kan District, Kandy, 2100', Uddawatakele Sane, 10-23.1.1970 (Davis & Rowe) 
(USNM, Washington); 1 $, Keg District, Lavant Estate, nr Yatiyantota, 80', 
19.xi.1970 (Flint) (USNM, Washington); 1 <$, Maskeliya, ix.1905 (Alston) (genitalia 
slide no. 19748; BMNH). 

Tinissa torvella mysorensis subsp. n. 

(PI. 16, fig. 96) 

Genitalia <£ unknown. 

Genitalia 9 (PI- 1 ^>. n g- 96)- Similar to torvella torvella but posterior projection of eighth 
sternite not as elongate, dorsally setose, not extending beyond tips of eighth tergite. Ostium 
situated slightly more ventrally; antrum almost three times as wide as in nominate sub- 
species and longer; bulbous and lightly sclerotized portion of antrum at point of junction 
with ductus seminalis larger than in torvella torvella and with three shallow lateral folds, wall 
deeply invaginated to form internal process partly blocking antrum immediately posterior 
to junction with ductus seminalis. 

Remarks. Differentiated from T. torvella torvella on characteristics of the $ 
genitalia as above, T. torvella mysorensis is not separable from the nominate sub- 
species on external characteristics. 

The holotype has a wing-span of 27 mm. 

Distribution. India (Mysore). 

Material examined. 

Holotype $, India: Mysore Prov., W. Ghats, Balehonnur, Netraconda, 2800- 
3800', in bungalow, prob. at lights (Evetts) (genitalia slide no. 19755; BMNH). 

Tinissa polystacta (Meyrick) 

(PI. 6, fig. 40; PL 16, fig. 97) 

Scardia polystacta Meyrick, 1918, Ann. Transv. Mus. 6 : 47. Lectotype $, South Africa: 
Natal, New Hanover, iii. 1913 (Hardenberg) (genitalia slide no. 8113 [Vari]; TM, Pretoria) 
[not examined]. 



REVISION OF THE TINISSINAE 295 

Polymnestra perilithias Meyrick. 1927, Exot. Microlepidopt. 3 : 331. Holotype <§, South 

Africa: Transvaal, Woodbush, n.i.1925 (Janse) (genitalia slide no. 8044 [Vari]; TM, Pretoria) 

[not examined] [synonymized by Gozmany & Vari, 1973 : 86]. 
Polymnestra perilithias Meyrick; Janse, 1968 : 91, pi. 50, figs 3, 4, pi. 101, figs 9-11 [redescrip- 

tion]. 
Scardia polystacta Meyrick; Janse, 1968 : 98, pi. 71, figs 3, 4, pi. 105, figs 5-8, pi. 118, fig. 9 

[redescription ; designation of lectotype]. 
Tinissa polystacta (Meyrick) ; Gozmany & Vari, 1973 : 86, figs 227, 228 [redescription; synonymy; 

new combination]. 

$. 11-5 mm. Similarly patterned to §; antennal cilia at least double width of flagellar 
segments. Legs whitish, foreleg as in $• Mid tibia with two broad, oblique brown bands on 
outer face; tarsus whitish, brown medial spot above on first segment, brown spot above at 
tips of other segments. Hind leg damaged but tibial tuft greyish. All spurs with brownish 
ring below apex. 

§. 12 mm. Head whitish, few brown scales laterally. Maxillary palpus five-segmented, 
folded. Haustellum absent. Labial palpus whitish, flecked with brown on outer surface, 
terminal segment with brownish subapical band. Antenna ochreous-whitish flecked with 
brown, first flagellar segment dark brown above, cilia one-half width of flagellar segments. 
Thorax and tegula brown, white posteriorly. Fore wing blackish brown mixed with diffuse 
orange-brown spots, patterned with conspicuous white spots at margins; fringe dark brownish 
grey. Hind wing dull brownish grey; fringe dull grey. Only left fore leg remaining, dull 
ochreous, greyish brown above but banded with whitish at articulations, fifth tarsal segment 
white. 

Genitalia <$ (Janse, 1968: pi. 50, fig. 4). Corema absent from eighth segment. Saccus 
rather short, triangular. Uncus lobes separated by broad membrane, slightly waisted at 
one-half, densely spined on internal surface towards tip. Subscaphium broad posteriorly 
with slight medial emargination; rod-like anteriorly. Juxta bifid, projecting posteriorly and 
hardly displaced dorsally, tips elongate, almost reaching tips of valves. Valve broad basally, 
elongate, tapering posteriorly, tip hooked slightly inward. Process from membrane between 
valve and juxta blade-like, reaching three-quarters length of valve. Labis very large, project- 
ing posteriorly, cowl-shaped, reaching tips of uncus. Aedeagus curved, moderately stout, 
tapered to sharp point from three-quarters. 

Genitalia $ (PI. 16, fig. 97). Eighth sternite short, almost square, two groups of setae 
at posterior margin; ostium at right angles to plane of sternite. Antrum two-thirds length 
of sternite, only thinly sclerotized at one-half at point of junction with ductus seminalis. Ductus 
bursae short, regular constrictions fine; ventral surface of ductus with finely rugose sclerotized 
plate anteriorly, plate 1 -5 times length of antrum. Eighth tergite posteriorly rounded, irregular 
row of setae on posterior margin. 

Remarks. T. polystacta is the smallest Tinissa species. The $ differs from all 
other Tinissa species in the ventral sclerotization of the ductus bursae. Males 
show affinities to T. spaniastra, possessing a similarly cowl- or hood-shaped labis 
(or fused pair of labides). The ventral positioning and deep bifurcation of the 
juxta in spaniastra is similar to that in polystacta, but the valves of the latter species 
are much more elongate. The tips of the uncus in polystacta are not hooked as in 
spaniastra. The bursa copulatrix is asignate: the 'minute phylliform signum' of 
Gozmany & Vari (1973 : 87) is a foreign body, apparently a scale. T. polystacta is 
also allied to T. cnltellata, the only other Tinissa species with a five-segmented 
maxillary palpus - see 'Remarks' for cultellata. 

Distribution. South Africa. 



296 g. s. robinson 

Material examined. 

South Africa: Natal, i $, Isputeni, 23.1.1916 (Janse) (genitalia slide no. 10318; 
BMNH) [paralectotype of polystacta]; 1 J, Umroti, iv. 1892 {Hutchinson) (genitalia 
slide no. 12294; BMNH). 



Tinissa cultellata (Gozmany & Vari) comb. n. 
(PL 6, figs 41, 42; PL io, fig. 64) 

Leptozancla cultellata Gozmany & Vari, 1973, Transv. Mus. Mem. 18 : 87, fig. 230. Holotype 
<3\ Uganda: Ruwenzori Range, Mahoma River, 6700', 13-16. viii. 1952 (Fletcher) (genitalia 
slide no. 12022 [Gozmany - remounted by Robinson]; BMNH) [examined]. 

<J. 17, 19 mm. Head yellowish cream, few brown scales near eyes. Maxillary palpus 
short, five-segmented, folded. Labial palpus yellowish cream, first and second segment heavily 
marked with blackish brown on outer surface, terminal segment with diffuse light brown flecks 
in basal half of outer surface. Antenna, including scape, blackish brown. Thorax yellowish 
cream, tegula white. Fore wing blackish brown [badly rubbed in both examples] with cream 
spots at costa and termen, traces of fine cream transverse striae, light dot at end of cell ; fringe 
blackish brown, cream where marginal spots are extended. Hind wing slightly ochreous white, 
flecked with pale grey anteriorly and towards termen; grey spots at termen extending into 
whitish fringe. Legs ochreous, flecked with blackish brown, fore leg particularly heavily 
marked above, hind leg hardly marked. Hind tibial tuft ochreous-whitish, scales sparse [rub- 
bed?]. All spurs with greyish brown streak below apex. 

9- [Wings detached; not in condition suitable for description.] 

Genitalia $ (PI. 10, fig. 64). Corema present in eighth segment. Saccus broad, almost 
square. Lobes of uncus quite widely separated, fully fused with vinculum, tapered, sharply 
pointed and heavily sclerotized; small circular basal process, long spine-like lateral process 
parallel to uncus lobe, small thorn-like lateral process at three-quarters. Subscaphium broad, 
not heavily sclerotized. Juxta, if present, not separable from valve which is large, heavily 
sclerotized, with four processes from steep internal surface, all almost reaching tip of valve 
which is slightly curved inward. Transtilla narrow; labides a pair of appressed lobate processes 
dorsal to aedeagus [description of transtilla and labides doubtful owing to damage of prepara- 
tion]. Aedeagus curved, short, stout, tapered from three-quarters. 

Genitalia 9- Eighth sternite narrow, lightly sclerotized medially, U-shaped posterior 
medial emargination, line of setae at posterior margin; about eight setae scattered from one- 
third to posterior margin on each side. Ostium small, anterior, pouch-like ; antrum constricted 
at three-quarters posteriorly, only lightly sclerotized; narrow hemi-cylindrical dorsal sclero- 
tized region on ductus bursae one-quarter length of antrum (see 'Remarks'). Ductus bursae 
with fine regular constrictions overlaid with fine pimpling, appearing rugose. Corpus bursae 
double length of ovipositor, narrow and tapering posteriorly. Eighth tergite rather square 
with line of setae at posterior margin, conspicuous heavily sclerotized ridge arched posteriorly 
from anterior corners ; anterior to ridge a series of six parallel transverse ridges ; anterior margin 
reflected posteriorly to form shallow pouch. 

Remarks. This is a bizarre species which differs from all other Tinissa species 
in several features: like T. polystacta it has a five-segmented maxillary palp but is 
much larger. It lacks the distinctive oblique dark bands on the outer surface of 
the mid tibia and the tarsal spotting common in other Tinissa species. Males are 
recognizable by their sharp uncus lobes with parallel lateral processes and the 
cluster of sharp peaks of the valval processes. T. cultellata is the only Tinissa 
species in which the juxta cannot be visually separated from the valve and may 



REVISION OF THE TINISSINAE 



297 



not even be present. Females are characterized by the very small anterior ostium 
and short antrum and ductus bursae. Ventral sclerotization of the ductus bursae 
(see below) and the ridging of the eighth tergite are peculiar to this species. Affini- 
ties are vague and are to the other divergent African species, T. polystacta (five- 
segmented maxillary palpus, shape of aedeagus and uncus) and T. poliophasma 
(rugose regular constrictions of the ductus bursae). The ventral sclerotization of 
the ductus bursae of polystacta may be homologous with the dorsal sclerotization 
of cultellata; in cultellata sclerotization is anterior to the point of junction with the 
ductus seminalis and is thus not a colliculum although it bears a striking resemblance 
to one. Both male genitalia preparations of this species are in poor condition. 

Distribution. Uganda (Ruwenzori Range). 

Material examined. 

Uganda: i $, 1 $, Ruwenzori Range, Mahoma River, 6700', 13-16. viii. 1952 
(Fletcher) (genitalia slide nos. 8462 [Gozmany], 12115 [Robinson]; BMNH). 

LEPTOZANCLA Meyrick 

Leptozancla Meyrick, 1920, Voyage Ch. Alluaud R. Jeannel Afr. or. 2, Microlepid. : 107. Type- 
species: Leptozancla talaroscia Meyrick, 1920, ibidem : 108, by original designation and 
monotypy. 

Leptozancla Meyrick ; Fletcher, 1929 : 125 [type-species]. 

Philagrias Meyrick, 1932, Trans, ent. Soc. Lond. 80 : 119. Type-species: Philagrias zelotica 
Meyrick, 1932, ibidem 80 : 119, by monotypy. [Synonymized by Gozmany & Vari, 1973 : 87.] 

Leptozancla Meyrick; Gozmany & Vari, 1973 : 87 [redescription]. 

Presence of mandible uncertain - head preparation not made owing to lack of available 
material. Maxillary palpus five-segmented, folded. Antenna extending to, or almost to, 
tip of fore wing, with dense whorl of cilia on each segment, cilia length equal to width of segment 
(L. talaroscia) or without cilia (L. zelotica). Fore wing narrow, apex oblique (talaroscia) or 
shorter, broader, resembling Tinissa (zelotica), 8-5-12-0 mm long; M present in cell; i? 5 just 
posterior to apex; R t and i? 5 separate. No elongate oval patch of small, flat, ovate scales 
on under surface of fore wing between A l+2 and posterior margin and no opposing rough scale 
patch on hind wing. Hind wing with M present in cell. Hind tibia with broad, rough, 
appressed scales, few sparse raised scales proximally (hind tibia of talaroscia unknown). 

Genitalia g. Corema of similar type to Tinissa present in eighth segment but reduced 
and without hairs in L. talaroscia. Saccus large, triangular or rounded apically. Vinculum 
broad, exceptionally so in L. zelotica. Uncus fused with vinculum in zelotica, narrow mem- 
branous zone between the two in talaroscia. Uncus lobes connected by sclerotized bridge in 
zelotica. Juxta large, well developed, with distinct ventral suture or sulcus in zelotica. Valve 
absent. No equivalent to process arising from membrane between valve and juxta as present 
in most Tinissa species. Labides and transtilla highly developed, labides forming elongate, 
posteriorly directed spines. Subscaphium narrow, short (talaroscia) or broad and elongate 
(zelotica). Aedeagus short and rather broad, exceptionally so in zelotica. 

Genitalia <j>. Unknown. 

Leptozancla talaroscia Meyrick 
(PI. 6, fig. 43; PI. io, fig. 65) 
Leptozancla talaroscia Meyrick, 1920, Voyage Ch. Alluaud R. Jeannel Afr. or. 2, Microlepid. : 108. 
Lectotype <$, Kenya: Mt Kenya, alpine grassland and tree-heath, 3300-3500 m, i-ii. 1912 
(Alluaud & Jeannel) (genitalia slide; MNHN, Paris) [not examined]. 



298 G. S. ROBINSON 

Leptozancla talaroscia Meyrick; Viette, 1951 : 83 [lectotype designated]. 
Leptozancla talaroscia Meyrick; Gozmany & Vari, 1973 : 87, fig. 229 [redescription] . 

o*. 25 mm. Head whitish, dark brown near eyes. Labial palpus whitish, first and second 
segments flecked with brown on outer face. Antenna as long as forewing, light brown, first 
flagellar segment darker, tip of scape whitish, pecten dark brown ; cilia dense, as long as width 
of flagellar segments. Thorax and tegula dark brown, posterior margins white. Fore wing 
[rubbed] dark greyish brown, traces of whitish spots at costa; fringe mixed whitish and greyish. 
Hind wing pale grey; fringe white. Legs rubbed and damaged; fore and mid legs ochreous- 
whitish with few sparse brownish scales above. 

$ unknown. 

Genitalia g (PI. 10, fig. 65). Corema absent from eighth segment but shallow pocket 
with thin, sclerotized bar at base present in tergosternal membrane. Saccus broad, rounded. 
Uncus fused with vinculum, short and broad, rounded lateral projections at four-fifths. Juxta 
very large, elongate, projecting posteriorly, with deep rounded posterior emargination ; dorsally 
with pair of curved, stout processes [one with tip broken in illustrated preparation]. Trans- 
tilla fused with dorsum of juxta and supporting labides, four elongate, posteriorly directed 
spines, central pair supporting sclerotized strip reflected anteriorly; subscaphium short, not 
connecting with anterior extension of labides. Aedeagus short, stout. 

Remarks. L. talaroscia is a larger and narrower-winged species than L. zelotica 
in which the juxta has a distinct medial ventral suture and a pair of small finger-like 
processes at the tip and a pair of lateral flaps. The labides are in the form of only 
a pair of posteriorly directed spines in zelotica. 

Distribution. Kenya (Mt Kenya). 

Material examined. 

Paralectotype <$, Kenya: Mt Kenya, 11500', i. 1912 (Alluaud & Jeannel) 
(genitalia slide no. 121 16 [Robinson]; BMNH). 



Leptozancla zelotica (Meyrick) 
(PL 6, fig. 44; PI. ii, fig. 66) 

Philagrias zelotica Meyrick, 1932, Trans, ent. Soc. Lond. 80 : 119. Holotype <$, Ethiopia: 
Djem-Djem Forest, c. 8000', 5-7. x. 1926 (Scott) (genitalia slide no. 10177 [Gozmany]; BMNH) 
[examined]. 

Leptozancla zelotica (Meyrick) Gozmany & Vari, 1973 : 88, fig. 231 [new combination, redescrip- 
tion] . 

<§. 18 mm. Head [very rubbed] whitish, brown scales around eyes. Labial palpus ochreous- 
whitish, blackish brown above and on outer surface of first and second segments, terminal 
segment with slight brownish flecks on outer surface. Antenna, including scape, ochreous, 
two-thirds length of forewing, cilia absent, pecten brownish. Thorax and tegula brownish, 
tegula white posteriorly. Fore wing badly rubbed, some blackish brown scales remaining, 
traces of yellowish cream spots at costa; fringe mixed grey and whitish. Hind wing light 
brownish grey; fringe whitish. Legs ochreous flecked with brownish above. Mid tibia with 
two very broad oblique dark brown bands on outer face, first to third tarsal segments with 
diffuse brown spot above. Hind tibia [Prubbed] with broad, ochreous appressed scales above, 
few protuberant scales towards base; second to fourth tarsal segments with diffuse brownish 
spot above. 

$ unknown. 



REVISION OF THE TINISSINAE 299 

Genitalia <$ (PI. 11, fig. 66). Corema present in eighth segment. Saccus triangular, 
vinculum very broad dorsally. Uncus lobes short, fused with vinculum, joined to each other 
by narrow dorsal sclerotized bridge. Subscaphium broad. Juxta with medial suture ventrally, 
elongate, distal half with two broad lateral flaps; tip of juxta terminating in two slightly curved 
finger-like processes. Dorsal surface of juxta with two heavily sclerotized pouch-like recesses. 
Labides fused with juxta via transtilla; reflected dorsally and anteriorly, then tapering pos- 
teriorly as elongate spines with broad, spur-like processes at one half. Aedeagus short and 
stout, extensive zone of microtrichia at tip. 

Remarks. The antennae of zelotica are shorter than in L. talaroscia, not thickly 
ciliate; the wings are narrower and less elongate. The structure of the juxta and 
labides separates the two species and talaroscia does not have a sclerotized 'bridge' 
between the uncus lobes. 

Distribution. Ethiopia. 

Material examined. 
Holotype only. 



references 

Diakonoff, A. 1952. Microlepidoptera of New Guinea. Part 1. Verh. K. ned. A had. 
Wet. (2) 49(i) : 1-167. 

1955- Microlepidoptera of New Guinea. Part 5. Verh. K. ned. Akad. Wet. (2)50(3) : 

1-210. 

1967. Microlepidoptera of the Philippine Islands. Bull. U.S. natn. Mus. 257 : 1-484. 

Fletcher, T. B. 1929. A list of the generic names used for Microlepidoptera. Mem. Dep. 

Agric. India (Ent.) 11 : i-ix, 1-244. 
Gozmany, L. A. 1969. The scientific results of Hungarian Zoological Expeditions to Tan- 
ganyika. No. 13. Tineid moths (Lepidoptera) collected by Dr J. Szunyoghy in the Mt. 

Meru area in 1965/66. Annls. hisl.-nat. Mus. natn. hung. 61 : 279-294. 

& Vari, L. 1973. The Tineidae of the Ethiopian region. Transv. Mus. Mem. 18 : 1-238. 

Janse, A. J. T. 1968. On the types of South African Microlepidoptera. Transv. Mus. Mem. 

16 : 1-127. 
Klots, A. B. 1956. Lepidoptera. In Tuxen, S. L. (Ed.), Taxonomist's glossary of genitalia 

in insects. Pp. 97-1 11, text-figs 121-132. Copenhagen. 
Meyrick, E. 1920. In Alluaud, Ch. & Jeannel, R., Voyage de Ch. Alluaud et R. Jeannel 

en Afrique Orientate (ign-igi2). Resultats scientifiques. Insectes Lepidopteres. 2. 

Microlepidoptera. Pp. 35-120. Paris. 

1928. Exot. Microlepidopt. 3 : 417-448. 

Viette, P. 195 1. Les types de Tineides de Meyrick appartenant au Museum de Paris. 

Bull. Soc. ent. Fr. 56 : 81-90. 
Walker, F. 1864. List of the specimens of the lepidopterous insects in the collections of the 

British Museum. 29. Tineites. Pp. iv -f 564-835. London. 
Zagulajev, A. K. 1972. New and little-known species of the moth families Tineidae, Deutero- 

tineidae and Ochsenheimeriidae (Lepidoptera). [In Russian.] Trudy zool. Inst. Leningr. 

52 : 332-356- 



3°o 



G. S. ROBINSON 

INDEX 

Synonyms are in italics; principal references are in bold. 



albipuncta sp. n., 264, 266, 274, 275 
amboinensis sp. n., 259, 264, 265, 267, 268, 

269, 273 
araucariae sp. n., 266, 273, 274, 275 

bakeri sp. n., 264, 281, 283, 284, 285, 292 
baliomicta Meyrick, 263, 264, 265, 270, 271, 
277, 278, 279, 280, 281, 282, 283, 284 

chalcites sp. n., 264, 266, 273, 274, 275 
chaotica sp. n., 257, 261, 263, 266, 271 
chloroplocama Meyrick, 273, 278, 285, 286 
cinerascens Meyrick, 259, 264, 265, 267, 268, 

269, 270, 271, 272 
convoluta sp. n., 266, 272 
cultellata Gozmany & Vari, 261, 262, 264, 

266, 267, 295, 2%, 297 

distracta Meyrick, 259, 264, 265, 268, 269, 

271, 272 
dohertyi sp. n., 265, 279, 284 

errantia sp. n., 264, 270, 272, 281 
eumetrota Meyrick, 266, 275 

Gelechia Hiibner, 283 

goliath sp. n., 265, 286, 287, 288 

heterograpta Meyrick, 277, 278, 279, 280 

indica sp. n., 257, 264, 265, 282, 284, 285 
insignis Zagulajev, 266, 279, 288 
insularia sp. n., 257, 263, 264, 265, 285, 286, 
287 

kidukaroka sp. n., 265, 279, 287, 288 
krakatoasp. n., 257, 259, 265, 267 



Leptozancla Meyrick, 255, 257, 259, 261, 263, 
296, 297 

mysorensis subsp. n., 265, 293, 294 

palmodes Meyrick, 263, 266, 272, 276 
parallela sp. n., 264, 266, 278, 279, 281, 282, 

286, 288 
perilithias Meyrick, 266, 295 
Philagrias Meyrick, 297, 298 
philippinensis sp. n., 266, 276, 277, 281 
phrictodes Meyrick, 257, 261, 264, 265, 289 
poliophasma Bradley, 261, 265, 266, 290, 

291, 297 
Polymnestra Meyrick, 266, 295 
polysema Zagulajev, 266, 276, 277 
polystacta Meyrick, 257, 261, 263, 264, 266, 

267, 289, 294, 295, 296, 297 

rigida Meyrick, 257, 264, 265, 277, 278, 279, 

280, 281, 282, 283 
ruwenzorica Gozmany & Vari, 259, 261, 263, 

266, 290, 292, 293 

Scardia Treitschke, 294, 295 
spaniastra Meyrick, 260, 261, 264, 265, 266, 
291, 295 

talaroscia Meyrick, 257, 261, 263, 297, 298, 

299 
Tinissa Walker, 255, 257, 261, 263, 266 
Tinissinae Gozmany & Vari, 255, 261 
torvella mysorensis subsp. n., 265, 293, 294 
torvella torvella Walker, 262, 265, 293, 294 
torvella Walker, 257, 259, 260, 261, 262, 263, 

266, 285, 292 
transversella Walker, 265, 283, 284 

zelotica Meyrick, 261, 263, 297, 298, 299 



G. S. Robinson, Ph.D. 

Department of Entomology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 



PLATE i 



Fig. i. Tinissa torvella Walker, (J, Sri Lanka, distal region of antenna, X 250. 

Fig. 2. T. torvella Walker, 2, Sri Lanka, antenna, X 550. 

Fig. 3. T. torvella Walker, 2, Sri Lanka, specialized scale area on forewing underside, X 80. 

Fig. 4. T. torvella Walker, 2, Sri Lanka, specialized scale, x 800. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 1 





PLATE 2 



Fig. 


5- 


Fig. 


6. 


Fig. 


7- 


Fig. 


8 


Fig. 


9- 


Fig. 


IO 


Fig. 


ii 


Fig. 


12 



Tinissa krakatoa sp. n., 9 holotype, Rakata I. [image reversed], 

T. cinerascens Meyrick, $ paralectotype, Sudest I. 

T. cinerascens Meyrick, § lectotype, Rossel I. 

T. distracta Meyrick, £ paralectotype, Assam. 

T. distracta Meyrick, £ lectotype, Assam. 

T. errantia sp. n.,^ holotype, Philippines. 

T. amboinensis sp. n., § holotype, Ambon I. 

T. amboinensis sp. n. )( J paratype, Ambon I. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 2 



/ 


£p 




-§$^~ 




6 







PLATE 3 



Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 19 
Fig. 20 



13- 

14. 

15- 
16. 

J 7- 
if 



Tinissa chaotica sp. n., $ holotype, Ambon I. 
T. convoluta sp. n., 9 holotype, Bougainville. 
T. palmodes Meyrick, 9 holotype, New Guinea. 
T. chalcites sp. n., 9 holotype, New Guinea. 
T. albipuncta sp. n., 9 holotype, New Guinea. 
T. eumetrota Meyrick, 9 lectotype, New Ireland. 
T. polysema Zagulajev, 9 holotype, Java. 
T. philippinensis sp. n., 9 holotype, Philippines. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 3 





PLATE 4 

Fig. 21. Tinissa rigida Meyrick, § [lectotype of heterograpta], New Britain. 

Fig. 22. T. rigida Meyrick, ^ holotype, Kai Is. 

Fig. 23. T. baliomicta Meyrick, § holotype, Philippines. 

Fig. 24. T. parallela sp. n. )C £ paratype, Sumatra. 

Fig. 25. 7\ parallela sp. n., 9 holotype, Sumatra. 

Fig. 26. 7\ indica sp. n., § holotype, Sikkim. 

Fig. 27. 7\ transvev sella (Walker), §, Sulawesi. 

Fig. 28. 7\ dohertyi sp. n., 9 holotype, Ambon I. 



Bull. Br. Mus. rial. Hist. (Ent.) 32, 7 



PLATE 4 




v 


» PBh89t 


^P 




.biG. 


29 


Fig. 


30 


Fig. 


3i- 


Fig. 


32 


Fig. 


33 


Fig. 


34 


Fig. 


35 


Fig. 


36. 



PLATE 5 

Tinissa insularia sp. n., 9 paratype, Java. 

T. insularia sp. n. )( J paratype, Penang. 

T. goliath sp. n., 9 holotype, New Guinea. 

T. kidukavoka sp. n., 9 holotype, Borneo. 

T. insignis Zagulajev, 9 holotype, Java. 

T. phrictodes Meyrick, <J paralectotype, Solomon Is. 

T. phrictodes Meyrick, 9 lectotype, Solomon Is. 

T. poliophasma Bradley, 9 holotype, Uganda. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 5 






PLATE 6 



Fig 
Fig 
Fig 
Fig 
Fig 
Fig 
Fig. 43 
Fig. 44 



37- 
38. 
39- 
40. 
41. 
42. 



Tinissa spaniastra Meyrick, $ holotype, Ethiopia [reversed image]. 

T. rnwenzorica Gozmany, $ holotype, Uganda. 

T. torvella torvella Walker, $, Sri Lanka [reversed image]. 

T. polystacta (Meyrick), Q paralectotype, South Africa. 

T. cultellata (Gozmany & Vari) )( J holotype, Uganda. 

T. cultellata (Gozmany & Vari),<J, Uganda. 

Leptozancla talaroscia Meyrick, $ paralectotype, Kenya. 

L. zelotica (Meyrick),^ holotype, Ethiopia. 



Bull. Br. Mus. nat. Hist. (Ent.) $>, 7 



['LATE 5 





PLATE 7 

Male genitalia 

Fig. 45. Tinissa distracta Meyrick, paralectotype, Assam. 

Fig. 46. T. cinerascens Meyrick, New Guinea. 

Fig. 47. T. cinerascens Meyrick, New Guinea (coremata). 

Fig. 48. T. amboinensis sp. n., paratype, Ambon I. 

Fig. 49. T. errantia sp. n., holotype, Philippines. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 7 




45 14 










\/f 



1 V 



47 




3 



If! 




Kr 



49 



rl 



PLATE 8 

Male genitalia 

Fig. 50. Tinissa chaotica sp. n., holotype, Ambon I. 

Fig. 51. T. chalcites sp. n., paratype, New Guinea. 

Fig. 52. T. albiptincta sp. n., paratype, New Guinea. 

Fig. 53. T. rigida Meyrick, holotype, Kai Is. 



Bull. By. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 8 





. ■*/ 




\1 







53 




PLATE 9 

Male genitalia 

Fig. 54. Tinissa parallela sp. n., paratype, Sumatra. 

Fig. 55. T. indica sp. n., paratype, Assam. 

Fig. 56. T. bakeri sp. n., holotype, Philippines. 

Fig. 57. T. insularia sp. n., paratype, Penang. 

Fig. 58. T. phrictodes Meyrick, Solomon Is. 

Fig. 59. T. spaniastra Meyrick, holotype, Ethiopia. 



Bull. Br. Mas. nat. Hist. (Ent.) 32, 7 



PLATE 9 







. &i 










56 



! f fiqP- 






55 







\ 






58 V 




" f 



>^- ■ ■ |l " " 



59 



Fig. 


60. 


Fig. 


61. 


Fig. 


62. 


Fig. 


63- 


Fig. 


64. 


Fig. 


65- 



PLATE 10 

Male genitalia 

Tinissa ruwenzorica Gozmany, holotype, Uganda. 
T. ruwenzorica Gozmany, holotype (eighth sternite). 
T. torvella torvella Walker, Sri Lanka. 
T. torvella torvella Walker, Sri Lanka (eighth sternite) . 
T. cultellata (Gozmany & Vari), holotype, Uganda. 
Leptozancla talaroscia Meyrick, paralectotype, Kenya. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 10 




60 






* 




64 



\ 







65 



M 



Fig. 


66. 


Fig. 


67. 


Fig. 


68. 


Fig. 


69. 


Fig. 


70. 



PLATE 11 

Male genitalia 

Leptozancla zelotica (Meyrick), holotype, Ethiopia. 

Female genitalia 

Tinissa krakatoa sp. n., holotype, Rakata I. 
T. cinerascens Meyrick, New Guinea. 
T. distracta Meyrick, lectotype, Assam. 
T. amboinensis sp. n., holotype, Ambon I. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 11 




66 







PLATE 12 



Fig. 


7i 


Fig. 


7- 


Fig. 


73 


Fig. 


74 


Fig. 


75 


Fig. 


76 


Fig. 


77 


Fig. 


78 


Fig. 


79 



Female genitalia 

Tinissa amboinensis sp. n., holotype (eighth tergite lobe). 

T. araucariae sp. n., holotype, New Guinea (eighth tergite lobe). 

T. chalcites sp. n., holotype, New Guinea (eighth tergite lobe). 

T. convoluta sp. n., holotype, Bougainville. 

T. eumetrota Meyrick, lectotype, New Ireland. 

T. araucariae sp. n., holotype, New Guinea. 

T. chalcites sp. n., holotype, New Guinea. 

T. albipuncta sp. n., holotype, New Guinea. 

T. palmodes Meyrick, holotype, New Guinea. 



Bull. Br. Mus. 11,1/. Hist. (Ent.) $2, 7 



PLATE 12 




Fig. 


80. 


Fig. 


81. 


Fig. 


82. 


Fig. 


83- 


Fig. 


84. 



PLATE 13 

Female genitalia 

Tinissa polysema Zagulajev, holotype, Java. 

T. philippinensis sp. n., holotype, Philippines. 

T. baliomicta Meyrick, New Guinea [see Text-fig. 6], 

T. rigida Meyrick [lectotype of heterograptd], New Britain [see Text-fig. 4]. 

T. baliomicta Meyrick, Philippines [see Text-fig. 5]. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 13 









PLATE 14 

Female genitalia 

Fig. 85. Tinissa parallela sp. n., holotype, Sumatra [see Text-fig. 7]. 

Fig. 86. T. indica sp. n., paratype, Bhutan. 

Fig. 87. T. insularia sp. n., paratype, Ambon I. 

Fig. 88. T. dohertyi sp. n., holotype, Ambon I. [see Text-fig. 8]. 

Fig. 89. T. transver sella (Walker), Sulawesi. 



Bull. Br. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 14 





86 




PLATE 15 

Female genitalia 

Fig. 90. Tinissa goliath sp. n., holotype, New Guinea. 

Fig. 91. T. kidukaroka sp. n., holotype, Borneo [see Text-fig. 9]. 

Fig. 92. T. insignis Zagulajev, holotype, Java [see Text-fig. 10]. 

Fig. 93. T. phrictodes Meyrick, New Ireland. 



Dull. By. Mus. nat. Hist. (Ent.) 32, 7 



PLATE 15 



' 




90 






PLATE i 6 

Female genitalia 

Fig. 94. Tinissa spaniastra Meyrick, Ethiopia. 

Fig. 95. T. torvella torvella Walker, Sri Lanka. 

Fig. 96. T. torvella mysorensis subsp. n., holotype, India. 

Fig. 97. T. polystacta (Meyrick), paralectotype, South Africa. 

Fig. 98. T. poliophasma Bradley, holotype, Uganda. 



Bull. Br. Mas. nat. Hist. (Ent.) 32, 7 



PLATE 16 



94 






97 




ENTOMOLOGY SUPPLEMENTS 



3. Watson, A. A revision of the Ethiopian Drepanidae (Lepidoptera). Pp. 177: 
18 plates, 270 text-figures. August, 1965. £4.20. 

4. Sands, W. A. A revision of the Termite Subfamily Nasutitermitinae (Isoptera, 
Termitidae) from the Ethiopian Region. Pp. 172: 500 text-figures. September, 

1965- £3-25. 

6. Okada, T. Diptera from Nepal. Cryptochaetidae, Diastatidae and Droso- 
philidae. Pp. 129: 328 text-figures. May, 1966. £3. 

7. Giliomee, J. H. Morphology and Taxonomy of Adult Males of the Family 
Coccidae (Homoptera: Coccoidea). Pp. 168: 43 text-figures. January, 1967. 

8. Fletcher, D. S. A revision of the Ethiopian species and a check list of the 
world species of Cleora (Lepidoptera: Geometridae) . Pp. 119: 14 plates, 146 
text-figures, 9 maps. February, 1967. £3.50. 

9. Hemming, A. F. The Generic Names of the Butterflies and their type-species 
(Lepidoptera: Rhopalocera) . Pp. 509. £8.50. Reprinted 1972. 

10. Stempffer, H. The Genera of the African Lycaenidae (Lepidoptera: Rho- 
palocera). Pp. 322: 348 text-figures. August, 1967. £8. 

11. Mound, L. A. A review of R. S. Bagnall's Thysanoptera Collections. Pp. 172: 
82 text-figures. May, 1968. £4. 

12. Watson, A. The Taxonomy of the Drepaninae represented in China, with 
an account of their world distribution. Pp. 151: 14 plates, 293 text-figures. 
November, 1968. £5. 

13. Afifi, S. A. Morphology and Taxonomy of Adult Males of the families 
Pseudococcidae and Eriococcidae (Homoptera: Coccoidea). Pp. 210: 52 text- 
figures. December, 1968. £5. 

14. Crosskey, R. W. A Re-classification of the Simuliidae (Diptera) of Africa and 
its Islands. Pp. 198: 1 plate, 331 text-figures. July, 1969. £4.75. 

15. Eliot, J. N. An analysis of the Eurasian and Australian Neptini (Lepidoptera: 
Nymphalidae). Pp. 155: 3 plates, 101 text-figures. September, 1969. £4. 

16. Graham, M. W. R. de V. The Pteromalidae of North-Western Europe 
(Hymenoptera: Chalcidoidea). Pp. 908: 686 text -figures. November, 1969. 

£19. 

17. Whalley, P. E. S. The Thyrididae of Africa and its Islands. Pp. 198: 

68 plates, 15 text-figures. October, 1971. £12. 

18. Sands, W. A. The Soldierless Termites of Africa (Isoptera: Termitidae). 
Pp. 244: 9 plates, 661 text-figures. July, 1972. £9.90. 

19. Crosskey, R. W. A Revisionary Classification of the Rutiliini (Diptera: 
Tachinidae), with keys to the described species. Pp. 167: 109 text-figures. 
February, 1973. £6.50. 

20. von Hayek, C. M. F. A Reclassification of the Subfamily Agrypninae 
(Coleoptera: Elateridae). Pp. 309: 17 text figures. October, 1973. £12.30. 

21. Crosskey, R. W. A Conspectus of the Tachinidae (Diptera) of Australia, 
including keys to the supraspecific taxa and taxonomic and host catalogues. 
Pp. 221: 95 text-figures. December, 1973. £9-55- 

printed by Unwin Brothers Limited the gresham press old woking surrey England 



_> +^l I . -p~ 



THE WESTERN PALAEARCTIC 

ICHNEUMONIDAE (HYMENOPTERA) 

OF BRITISH AUTHORS 




M. G. FITTON 



BULLETIN OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 
ENTOMOLOGY Vol. 32 No. 8 

LONDON : 1976 



THE WESTERN PALAEARCTIC 

ICHNEUMONIDAE (HYMENOPTERA) 

OF BRITISH AUTHORS 



BY 



MICHAEL GEOFFREY FITTON 



Pp. 301-373 




BULLETIN OF 

THE BRITISH MUSEUM (NATURAL HISTORY) 

ENTOMOLOGY Vol. 32 No. 8 

LONDON : 1976 



THE BULLETIN OF THE BRITISH MUSEUM 

(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they become 
ready. Volumes will contain about three or four 
hundred pages, and will not necessarily be completed 
within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 
Department. 

This paper is Vol. 32 No. 8 of the Entomology 
series. The abbreviated titles of periodicals cited 
follow those of the World List of Scientific Periodicals. 



World List abbreviation 
Bull. Br. Mus. nat. Hist. (Ent. 

ISSN 0524-6431 



Trustees of the British Museum (Natural History), 1976 



TRUSTEES OF 
THE BRITISH MUSEUM (NATURAL HISTORY) 

Issued 27 May, 1976 Price £4.95 

This issue completes volume 32. 



THE WESTERN PALAEARCTIC 

ICHNEUMONIDAE (HYMENOPTERA) 

OF BRITISH AUTHORS 





By M. G. FITTON 








CONTENTS 


Page 


Synopsis ...... . 304 


Introduction .... 


. 






304 


Format OF CATALOGUE ENTRIES. 








305 


Depositories of material 








306 


Catalogue of western Palaearctic 


Ichneumonidae 


of Britisi 


i 




authors ..... 








3°7 


BlGNELL, G. C. 










3°7 


Bridgman, J. B. 










308 


Bridgman, J. B. & 


: Fitch, E. A. 








320 


Cameron, P. 










321 


Capron, E. . 










321 


Curtis, J. 










322 


Desvignes, T. 










324 


FORSTER, J. R. 










331 


Haliday, A. H. 










331 


Kerrich, G. J. 










335 


KlRBY, W. 










336 


Laidlaw, W. B. R 










336 


Lewin, W. . 










337 


Marshall, T. A. 










337 


Morley, C. 










339 


Parfitt, E. . 










345 


Perkins, J. F. 










34 6 


Stelfox, A. W. 










348 


Stephens, J. F. 










349 


Walker, F. 










354 


Waterston, J. 










355 


Westwood, J. O. 








355 


WOLLASTON, T. V. 








355 


Summary of new combinations and 


NEW SYNONYMS 






35 6 


New combinations 








356 


New synonyms 


. 






357 


Summary of nomina dubia 








357 


List of replacement names 








358 


Acknowledgements 








358 


References .... 








358 


Index 




. 






364 



304 M. G. FITTON 

SYNOPSIS 

The 529 species-group names proposed by British authors for western Palaearctic Ichneu- 
monidae (Hymen optera) are catalogued. The generic placements of the species to which the 
names apply are established after study of the extant types. Types of 75 species are lost 
and 35 names remain nomina dubia. Lectotypes are designated for 144 nominal species- 
group taxa. Seventy-one new combinations and 13 new specific synonyms are established. 

INTRODUCTION 

This paper is intended as a contribution to a complete catalogue and reclassification 
of the western Palaearctic Ichneumonidae. Catalogues of this kind are essential 
if revisionary work is to have a sound nomenclatural basis. The main aims of 
the present work have been to list all of the relevant names proposed by British 
authors, locate type-material and establish the generic placements of the species 
after study of the types. 

The need to catalogue the western Palaearctic Ichneumonidae is all the more 
apparent now that the catalogues produced by H. and M. Townes and co-workers 
cover the rest of the world. The volume and complexity of the literature dealing 
with the western Palaearctic fauna make a complete bibliographic treatment of 
a similar kind virtually impossible. However, the basic data (on names, types 
and generic placements) that are essential prerequisites to revisionary studies 
can be collected and presented in a readily usable form fairly quickly. Catalogues 
of this kind treating the names of one or a group of authors are more easily checked 
for completeness than those covering groups defined taxonomically because of the 
very large number of ichneumonid names that have never been properly accounted 
for. Another advantage of this approach is that the names and type-material 
of any one author tend to be in a restricted number of publications and collections 
respectively. 

The western Palaearctic Ichneumonidae were last catalogued completely by 
Dalla Torre (1901; 1902). Morley made an attempt to account for all of the British 
species of British authors in his Ichneumonologia Britannica (1903-1915) but his 
work is unreliable and is now largely ignored. More recently Perkins (1953) has 
studied the British Ichneumoninae and Horstmann (1972) has catalogued a large 
proportion of the species described by Bridgman. Oehlke (1967) and Aubert 
(1969a) have produced catalogues of the western Palaearctic Pimplinae (Aubert's 
catalogue also includes the Xoridinae and Acaenitinae) . 

In this paper an attempt has been made to account for all of the species-group 
names proposed by British (including Irish) authors for western Palaearctic Ichneu- 
monidae up to July 1975. Nomina nuda have been omitted. For convenience 
the limits of the western Palaearctic have been taken as the western boundary of 
the eastern Palaearctic as defined by Townes, Momoi & Townes (1965) and the 
northern boundary of the Ethiopian region as defined by Townes & Townes (1973). 
A small number of species described from the western Palaearctic in error are 
included in the catalogue. 

A large proportion of the species dealt with in this catalogue were described 
before the practice of designating a single specimen as 'holotype' or 'type' was 



WESTERN PALAEARCTIC ICHNEUMONIDAE 305 

adopted. Indeed in many of the original descriptions there is little or no reference 
to the type-material. When only a single type-specimen has been found, and there 
is no evidence that more than one original specimen existed, that specimen has been 
accepted and is cited as the 'holotype'. The treatment of single extant types, 
from type-series of unknown size, as holotypes is discussed by Crosskey (1974 : 272- 
275) and is the practice followed in the Townes catalogues (e.g. Townes & Townes, 
J 973 : 4)- The more positive approach of designating such specimens as lectotypes, 
advocated by Vane-Wright (1975 : 26-28), has not been adopted. 

There are previous valid type restrictions for a number of the species originally 
described from syntype series and in addition a large number of lectotypes are 
designated in this paper. It has been thought desirable to designate lectotypes 
because of the past treatment of type-material, particularly in the BMNH collection. 
It was Morley's practice to select and label one specimen from a syntype series as 
'type'. There is evidence in some of Morley's statements that some 'types' had 
been labelled by earlier curators of the BMNH collection and others have been 
labelled subsequently by J. F. Perkins. Unfortunately most of these type selections 
have never been published and in a large number of cases it is not now possible 
(because of inadequate labelling) to identify other members of the original syntype 
series. Morley was very inconsistent when dealing with types in his published 
work although he clearly had a 'type' concept. These 'types' have been regarded 
in the same way as holotypes and in the BMNH were serially numbered and segre- 
gated in a 'Type collection' together with genuine holotypes and properly designated 
lectotypes. In cases of this kind the lectotypes designated here are the specimens 
that have been traditionally regarded as 'types' unless otherwise stated. 

The generic placements given in this paper mostly follow the work of Townes 
(1969; 1970a; 1970&; 1971). My inexperience means that some species will have 
been misplaced and this is likely to be the main shortcoming of the catalogue. 
J. F. Perkins' work with the BMNH collection has been invaluable as a guide to 
the placement of a large proportion of the species. In the cases where types have 
been lost or destroyed and there has been no recent reliable consideration of place- 
ment the names are given as nomina dnbia. Little attention has been paid to 
specific synonymy since the extra work involved would in any case need to be 
repeated during re visionary studies. Previously published synonyms are given 
where they are probably correct and some new synonyms are established where 
there can be little doubt about the identity of the species concerned. Limited 
bibliographic references are cited to papers dealing with the placements and synony- 
mies given. Usually the reference is to the most recent authoritative work, not 
necessarily to where the placement or synonymy was established. 

FORMAT OF CATALOGUE ENTRIES 

For each name the entry is arranged in the following sequence. 

Name; author; date and page reference of original publication; status and 
sex of primary type(s); locality of primary type(s); type-depository; lectotype 
designation or reference to previous valid type restriction (when necessary). 



3 o6 M. G. FITTON 

Number and sex of paralectotypes, with data as for lectotype (when necessary). 

Statements on the labelling of the type-material especially with regard to its 
identification as such and on its condition if badly damaged. 

A statement, prefixed 'Identity', on the generic placement and synonymy 
of the species. 

The following points should be noted with regard to these data. 

The name is given as published except that the orthography is altered as 
necessary to comply with Articles 26, 27, 28 and 32 of the International 
Code of Zoological Nomenclature (1961). 

The date given is the actual date of publication. If a different date is given 
in the original publication this is shown in the list of references and any relevant 
information is given in the notes on individual authors. Dates of publication 
of parts of the Trans, ent. Soc. Lond. are given by Wheeler (1912). 

In the cases of types lost or destroyed their status and sex is given on the 
basis of data available in the original description. 

Type-locality information not taken from the original description and/or 
specimen labels is enclosed in square brackets. English and Welsh county 
names in square brackets are the new counties operative from April, 1974. The 
names of the new Scottish regions have not been used because of their more 
recent introduction (May, 1975) and the very large areas which they cover. 

Where types have not been located 'lost' or 'destroyed' (on the evidence avail- 
able) is given in place of a type-depository. The abbreviations used to indicate 
type-depositories are given in the following section. 

Previous valid type restrictions. If the word lectotype was used in the restric- 
tion it is cited as 'by designation of . . .' or as 'by fixation of . . .' if the word 
lectotype was not used. 

The information on paralectotypes is omitted if it was given clearly and com- 
pletely with a previous lectotype designation. The phrase 'same data as lectotype' 
refers only to the locality data as given for the lectotype. 



DEPOSITORIES OF MATERIAL 

An annotated list of depositories, together with the abbreviations used to indicate 
them, is given below. Further notes on the collections of individual authors are 
given in the catalogue. 

BMNH British Museum (Natural History), London 

Contains the collections of Cameron, Capron, Desvignes, Fitch (part), 
Marshall, Morley, Stephens and Wollaston and some or all types of 
Bignell, Bridgman, Bridgman & Fitch, Cameron, Capron, Curtis, 
Desvignes, Kerrich, Laidlaw, Marshall, Morley, Perkins, Stelfox, Stephens 
Waterston and Wollaston. All material is incorporated into a single 
collection of world Ichneumonidae. Morley (1910 : 167-168 and 170) 
apparently amalgamated the main British collections and it was probably 
he who was responsible for labelling a large number of specimens as 
'types'. Many of these specimens came from syntype series which 



WESTERN PALAEARCTIC ICHNEUMONIDAE 



3°7 



CM, Norwich 



CMAG, Plymouth 



IRSNB, Brussels 
MHN, Geneva 
MNHN, Paris 
MNHU, Berlin 
NMI, Dublin 



NMV, Melbourne 



NR, Stockholm 
PEM, London 

UM, Oxford 
USNM, Washington 



UZI, Lund 
ZI, Wroclaw 
ZIU, Uppsala 
ZMU, Helsinki 
ZSBS, Munich 



cannot now be reconstructed as the original placements of specimens 
in the individual collections are not indicated (except for a proportion 
of the 'types' and a small number of specimens labelled as 'co-types'). 
Castle Museum, Norwich 

Contains the collection of Bridgman which includes most of Bridgman's 
types and types of Bridgman & Fitch and Parfitt. 
City Museum and Art Gallery, Plymouth 

Contains the collection of Bignell. The primary types, of Bignell, 
Bridgman and Marshall, from this collection are now on permanent 
loan at the BMNH. 

Institut Royal des Sciences Naturelles de Belgique, Brussels 
Museum d'Histoire Naturelle, Geneva 
Museum National d'Histoire Naturelle, Paris 
Museum fur Naturkunde der Humboldt-Universitat, Berlin 
National Museum of Ireland, Dublin 

Contains the collection of Haliday which includes the Haliday types 
and some Curtis types. 
National Museum of Victoria, Melbourne 

Contains the Curtis collection which includes most of the Curtis 
types and one Desvignes type. 
Naturhistoriska Riksmuseet, Stockholm 
Passmore Edwards Museum, London 

Contains part of the Fitch collection. 
University Museum, Oxford 
United States National Museum, Washington 

Contains the Stelfox collection which includes all except one of 
Stelfox's types. 

Universitetets Zoologiska Institution, Lund 
Zoological Institute, Wroclaw 
Zoological Institute of the University, Uppsala 
Zoological Museum of the University, Helsinki 
Zoologische Sammlung des Bayerischen Staates, Munich 



CATALOGUE OF WESTERN PALAEARCTIC ICHNEUMONIDAE OF 

BRITISH AUTHORS 

The catalogue is arranged in alphabetical order of authors and original binominal 
combinations. 



Bignell, G. C. 

Bignell's collection is in the City Museum and Art Gallery, Plymouth. The 
specimens are usually labelled with a printed number, the complete data being 
in a manuscript notebook kept with the collection. The ichneumonid primary 
types from the collection have been placed on permanent loan at the British Museum 
(Natural History) . Hodgson (1917) published an account of type-material in the 
Bignell collection. From a study of the collection (where a number of specimens 
are incorrectly labelled 'type') and the paper it is clear that Hodgson did not intend 
to make type restrictions and his mentions of 'type' are not construed as such. 

Pimpla bridgmanii Bignell, 1894a : 280. Holotype ?> [England:] Devon, Cann Woods 
(BMNH, on permanent loan from CMAG, Plymouth) [examined]. 
The holotype lacks the tips of the antennae. 



308 M. G. FITTON 

Bignell (18946 : 255) published a second description of this species under the name bridgmani. 
Identity. Junior synonym of Dreisbachia pictifrons (Thomson) (Oehlke, 1967 : 20). 

Pimpla epeirae Bignell, 1893 : 37. LECTOTYPE ?, [England:] Devon, Ivybridge (BMNH), 
here designated [examined]. 

Paralectotypes. 2 $, same data as lectotype (CMAG, Plymouth). 

Identity. Junior synonym of Tromatobia variabilis (Holmgren) (Oehlke, 1967 : 18). 

Pimpla ruflpleura Bignell, 1889 : xv. LECTOTYPE $, [Great Britain] (BMNH, on 
permanent loan from CMAG, Plymouth), here designated [examined]. 

Paralectotypes. 2 6* (BMNH, on permanent loan from CMAG, Plymouth) ; 1 q* (CMAG, 
Plymouth). All same data as lectotype. 

The lectotype is on the same mount as two of the paralectotypes. 

Identity. Tromatobia rufipleura (Bignell) (Oehlke, 1967 : 18). 



Bridgman, J. B. 

Bridgman's collection is in the Castle Museum, Norwich. The arrangement 
of specimens does not seem to have been altered significantly since Bridgman 
donated the collection to the museum in 1895. This, together with the published 
and unpublished work of J. F. Perkins and K. Horstmann, has greatly facilitated 
cataloguing of the type-material. Generally the only data with specimens are a 
handwritten number and/or a colour-coded label. The complete data are in a 
manuscript notebook kept with the collection. There is also a further volume of 
Bridgman's manuscript notes. 

Anomalon cylindricum Bridgman, 1884 (November) : 424. LECTOTYPE $, [Great 
Britain] (PEM, London), here designated [examined]. 

Paralectotype $, same data as lectotype (PEM, London). 

The two syntypes were standing in the Fitch collection under the label '1419. insidiator, 
Fst.' cut from Morley's catalogue. 

The lectotype and paralectotype are the same specimens as the lectotype and paralectotype, 
respectively, of Anomalon cylindricum Bridgman & Fitch. 

Identity. Junior primary homonym and synonym of Anomalon cylindricum Bridgman & 
Fitch. Valid name Barylypa cylindricum (Bridgman & Fitch). 

Anomalon minutum Bridgman, 1884 (November) : 425. LECTOTYPE $, [England: 
Hampshire,] New Forest (CM, Norwich), here designated [examined]. 

Paralectotypes. 2 6\ 5 $> same data as lectotype (CM, Norwich). 

The lectotype and paralectotypes are the same specimens as the lectotype and paralecto- 
types, respectively, of Anomalon minutum Bridgman & Fitch. The specimen labelled as 
'type' by Morley (1915c : 254-255) was (according to Bridgman's notes) collected in 1885 
and therefore has no type status. 

Identity. Junior primary homonym and synonym of Anomalon minutum Bridgman & 
Fitch. Valid name Agrypon minutum (Bridgman & Fitch) (det. I. D. Gauld). 

Anomalon nigripes Bridgman, 1887 : 150. LECTOTYPE <$, [England:] Sussex, Abbots- 
wood (CM, Norwich), here designated [examined]. 

Paralectotype 6*, same data as lectotype (CM, Norwich). 
Identity. Agrypon nigripes (Bridgman) (det. I. D. Gauld). 

Apterophygus paradoxus Bridgman, 1889 : 417. Holotype $, [England: Wiltshire,] 
Nunton (BMNH) [examined]. 

Identity. Phygadeuon paradoxus (Bridgman) (Horstmann, 1972 : 223). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 309 

Aptesis foersteri Bridgman, 1882 : 146. Holotype $, [England:] Norfolk, Brundall (CM, 
Norwich) [examined]. 

Identity. Junior synonym of Catalytus mangeri (Gravenhorst) (Horstmann, 1972 : 219). 

Bassus abdominator Bridgman, 1886 : 365. Holotype $, [England:] near Plymouth 
(CM, Norwich) [examined]. 

Identity. Syrphoctonus abdominator (Bridgman) (Diller, 1969 : 550). 

Bassus holmgreni Bridgman, 1882 : 161. LECTOTYPE $, [England:] Norfolk, Brundall 
(CM, Norwich), here designated [examined]. 

Paralectotype <$, [England:] Norfolk, Felthorpe (CM, Norwich). 

Identity. Junior synonym of Tymmophorus rufiventris (Gravenhorst) (Dasch, 1964 : 82). 

Bassus punctatus Bridgman, 1888 : 375. Holotype $, [England: Surrey,] Caterham [not 
Aviemore as stated by Bridgman] (CM, Norwich) [examined]. 

The specimen agrees in all respects with the original description except the locality, which 
is written (in ? Champion's handwriting) on the underside of the mount. There can be 
little doubt that this is Bridgman's original specimen. 

Identity. Junior synonym of Homotvopus fissorins (Gravenhorst) (Thomson, 1890 : 1504). 

Bassus scutellaris Bridgman, 1886 : 364. Holotype $, [England:] near Plymouth (CM, 
Norwich) [examined]. 

Identity. Junior primary homonym of Bassus scutellaris Cresson, 1868 : 112. Replace- 
ment name here proposed: Promethes bridgmani nom. 11. 

Bassus tibialis Bridgman, 1883 : 170. Syntype9, [Great Britain] (CM, Norwich) [examined]. 
The only extant syntype is in poor condition, being covered by the remains of a growth 
of mould, and is not designated as lectotype for that reason. The specimen was standing, 
unlabelled, under tibialis in Bridgman's collection and is probably the one from Mr Norgate. 
The other two specimens were probably returned to Mr Fletcher and as yet have not been 
traced. 

Identity. Junior primary homonym of Bassus tibialis Cresson. Syn. n. of Bioblapsis 
polita (Vollenhoven, 1878 : 161 {Trichomastix)) , holotype [?] $, [Netherlands:] Scheven- 
ingen (? depository) [not examined]. 

Campoplex costulatus Bridgman, 1886 : 346. Holotype $ [Great Britain] (lost). 

This specimen is presumably also the type of Campoplex costulatus Bridgman & Fitch. 
Identity. Junior primary homonym and synonym of Campoplex costulatus Bridgman & 
Fitch. Valid name Dusona anceps (Holmgren) (Hinz, 1972 : 47). 

Campoplex Jemorator Bridgman, 1886 : 347. Lectotype $, [Great Britain] (CM, Norwich), 
by designation of Hinz (1972 : 47) [examined]. 

Paralectotypes. 1 <J (CM, Norwich); 2 <J, 2 $ (BMNH). All same data as lectotype. 

The paralectotypes in the BMNH collection are 1 <J, 2 $ from the Harwood collection 
labelled 'C. femorata named by Bridgman' and 1 $ from the Capron collection (originally 
pinned but remounted on card by Capron) given to him by Bridgman (Morley, 1915c : 77). 
All specimens except the one from the Capron collection are mounted on similar short brass 
pins. 

The lectotype and paralectotypes are the same specimens as the lectotype and paralecto- 
types, respectively, of Campoplex femorator Bridgman & Fitch. 

Identity. Junior primary homonym and synonym of Campoplex femorator Bridgman & 
Fitch. Valid name Dusona polita (Foerster) (Hinz, 1972 : 47). 

Campoplex incompletus Bridgman, 1889 : 420. Holotype <J, [England:] Eastbourne 
(CM, Norwich) [examined]. 

Identity. Dusona incompleta (Bridgman) (Hinz, 1972 : 47). 

Campoplex punctatus Bridgman, 1886 : 345. Holotype <$, [Great Britain] (CM, Norwich) 
[examined]. 
This specimen is also the holotype of Campoplex punctatus Bridgman & Fitch. 



310 M. G. FITTON 

Identity. Junior primary homonym and synonym of Campoplex punctatus Bridgman & 
Fitch. Valid name Dusona petiolator (Fabricius) (Hinz, 1972 : 47). 

Cecidonomus gallicola Bridgman, 18806 : 265. Lectotype?, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 218) [examined]. 

Paralectotypes. 2 q\ 3 ?, same data as lectotype (BMNH). 
Identity. Mastrus gallicola (Bridgman) (Horstmann, 1972 : 218). 

Cecidonomus rufus Bridgman, 18806 : 265. Lectotype $, [England:] near Norwich, 
Eaton (CM, Norwich), by designation of Horstmann (1972 : 219) [examined]. 
Paralectotypes. 2 $ (BMNH); 1 $ (CM, Norwich). 
Identity. Junior synonym of Mastrus inimicus (Gravenhorst) (Horstmann, 1972 : 219). 

Cecidonomus westoni Bridgman, 18806 : 264. Lectotype?, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 218) [examined]. 

Paralectotypes. 1 <J (CM, Norwich) ; 7 £, 8 $ (BMNH). All same data as lectotype. 
Identity. Mastrus westoni (Bridgman) (Horstmann, 1972 : 218). 

Chorinaeus flavipes Bridgman, 1881 : 165. Holotype <j>, [England:] Norwich, Mousehold 
(CM, Norwich) [examined]. 

The holotype has lost the tips of the fore tarsi. 
Identity. Chorinaeus flavipes Bridgman. 

Cryptus antennatus Bridgman, 1881 : 153. Lectotype $, [England: near Norwich,] Eaton 
(CM, Norwich), by designation of Horstmann (1972 : 219) [examined]. 
Paralectotype $, same data as lectotype (CM, Norwich). 
Identity. Junior synonym of Enclisis macilenta (Gravenhorst) (Horstmann, 1972 : 219). 

Dicoelotus cameroni Bridgman, 1881 : 146. Lectotype ?, [England:] Norwich, Heigham 
(CM, Norwich), by fixation of Perkins (1959 : 86) [examined]. Dicoelotus is a lapsus for 
Dicaelotus. 

? Paralectotype $, [Scotland] (BMNH). 

The specimen in the BMNH was labelled '?Type' by Morley and is from Cameron's collec- 
tion. It may be the second syntype. 

Identity. Dicaelotus cameroni Bridgman (Perkins, 1959 : 86). 

Euryproctus minutus Bridgman, 1886 : 358. LECTOTYPE <£, [England:] near Norwich, 
Brundall (CM, Norwich), here designated [examined]. 
Paralectotype o". same data as lectotype (CM, Norwich). 
Identity. Syntactus minutus (Bridgman) comb. n. 

Exochus fletcheri Bridgman, 1884 : 432. LECTOTYPE $, [England: Cambridgeshire,] 
Wicken Fen (CM, Norwich), here designated [examined]. 

Paralectotypes. 1 o\ 1 $ [not 2 ^ as stated by Bridgman], same data as lectotype (CM, 
Norwich) . 

Bridgman described the females as males and vice versa. 

Identity. Exochus fletcheri Bridgman. 

Exochus niger Bridgman, 1883 : 169. Holotype $, [England:] Norwich, Norkhouse Lane 

(CM, Norwich) [examined]. 

Identity. Triclistus niger (Bridgman) (Aeschlimann, 1973 : 243). 

Glypta annulata Bridgman, 1890a : 71. LECTOTYPE $, [England: West Sussex,] Slindon 
(CM, Norwich), here designated [examined]. 

Paralectotype $, same data as lectotype (CM, Norwich). 

Bridgman's formal description of this species (unpublished manuscript) gives the informa- 
tion that it was based on two females bred by W. H. B. Fletcher from Sericornis conchana 
from Slindon and that it resembles G. pictipes except that the coxae are black. The two 
syntypes were standing in Bridgman's collection under the label '?sp. like pictipes T. but 
coxae black' (now attached to the lectotype) and have the corresponding collection data. 

Identity. Glypta annulata Bridgman. 



WESTERN PALAEARCTIC ICHNEUMONID AE 



3ii 



Glypta parvicaudata Bridgman, 1889 : 435. LECTOTYPE $, [Scotland: Outer Hebrides,] 
Stornoway (CM, Norwich), here designated [examined]. 
Paralectotype $, same data as lectotype (CM, Norwich). 
Identity. Glypta parvicaudata Bridgman. 

Glypta parvicornuta Bridgman, 1886 : 367. Holotype $, [England:] Cambridgeshire 
(CM, Norwich) [examined]. 

The holotype lacks the tips of the antennae and hind tarsi. 
Identity. Glypta parvicornuta Bridgman. 

Glypta rubicunda Bridgman, 1890a : 70. LECTOTYPE $, [Great Britain] (CM, Norwich), 
here designated [examined]. 

Paralectotypes. 4 $, 2 $, same data as lectotype (CM, Norwich). 

The specimen selected as lectotype is the female bearing J. F. Aubert's det. label. 
Bridgman (1890& : 209) published a second, more formal, description of this species. 

Identity. Glypta rubicunda Bridgman. 

Glypta rufata Bridgman, 1888 : 378. LECTOTYPE $, [England: Cambridgeshire,] Wicken 
Fen (CM, Norwich), here designated [examined]. 

Paralectotypes. 3d 1 . 1 ?, same data as lectotype (CM, Norwich). 

The specimen selected as lectotype is the female labelled 'type' by J. F. Perkins and bearing 
J. F. Aubert's det. label. 

Identity. Glypta rufata Bridgman. 

Glypta similis Bridgman, 1886 : 367. LECTOTYPE $, [England :] Worthing (CM, Norwich), 
here designated [examined]. 

Paralectotypes. 3 <J, 1 ?, same data as lectotype (CM, Norwich). 

The specimen selected as lectotype is the female bearing J. F. Aubert's det. label. 

Identity. Glypta similis Bridgman. 

Glypta trochanterata Bridgman, 1886 : 368. LECTOTYPE $, [England:] Norwich, 
Mousehold (CM, Norwich), here designated [examined]. 

Paralectotypes. 4 <J, 2 $, same data as lectotype (CM, Norwich). 

The specimen selected as lectotype is the female labelled 'type' by J. F. Perkins and bearing 
J. F. Aubert's det. label. 

Identity. Glypta trochanterata Bridgman. 

Grypocentrus bipunctatus Bridgman, 1886 : 358. Holotype $, [England:] Surrey, Wimble- 
don (CM, Norwich) [examined]. 

Identity. Lathrolestes bipunctatus (Bridgman). Generic placement established by Morley 
(1911c : 278, as Lathrolestus) here confirmed. 

Hemimachus annulicortiis Bridgman, 1883 : 160. Syntypes 3 o*. [Wales: Dyfed,] Milford 
Haven (BMNH) [examined]. 

The syntypes were standing in the Fitch collection, together with females, under the 
label 'annulicornis m.s.' (in Marshall's handw : riting). 

Identity. Junior synonym of Gelis rufulus (Foerster) (Horstmann, 1972 : 221). 

Hemimachus confusus Bridgman, 1883 : 159. ? Syntypes 3 <J. [England: Devon,] Barn- 
staple (BMNH) [examined]; 1 £, [England: Surrey,] Box Hill (BMNH) [examined]. 

The four specimens were standing, together with twenty females and one other male (not 
a Marshall specimen), in the Fitch collection under the name ' ' rufocinctus , Gr.'. From other 
evidence (see under Hemimachus annulicornis Bridgman and Phygadeuon marshalli Bridgman) 
there can be little doubt that 'a collection that belonged to Mr Marshall' was in Fitch's 
possession and thus it seems likely that the four males referred to above are the syntypes 
of H. confusus. One is labelled 'Box Hill' on the underside of the mount and the remainder 
'B' (= Barnstaple (Morley, 1915a)). 

Identity. ? Gelis confusus (Bridgman) comb. n. 



3 i2 M. G. FITTON 

Hemimachus hyponomeutae Bridgman, 1883 : 155. ? Holotype o\ [Great Britain] (lost). 
Identity. Junior synonym of Gelis corruptor (Foerster) (Horstmann, 1972 : 221). There 
are two specimens labelled 'Bred from Hyponomeuta evonymellus. n.s. type' in the Bignell 
collection (CMAG, Plymouth). 

Hemimachus ovatus Bridgman, 1883 : 158. Holotype q\ [England: Norfolk,] Brundall 
(CM, Norwich) [examined]. 

Identity. Junior synonym of Gelis formicarius (Fabricius) (Horstmann, 1972 : 221). 

Hemimachus piceus Bridgman, 1883 : 153. Lectotype <J, [England:] near Norwich, 
Mousehold (CM, Norwich), by designation of Horstmann (1972 : 221) [examined]. 
Paralectotypes. 2 q\ same data as lectotype (CM, Norwich). 
Identity. ? Hemiteles piceus (Bridgman) (Horstmann, 1972 : 221). 

Hemimachus ruflpes Bridgman, 1883 : 157. Syntype o*, [England:] Norwich, Luckswood 
Lane (CM, Norwich) [examined]. 

Identity. ? Junior synonym of Gelis cantus (Foerster) (Horstmann, 1972 : 221). Gelis 
rufipes (Bridgman) is a junior secondary homonym of G. rufipes (Foerster). 

Hemimachus rufotinctus Bridgman, 1883 : 155. Holotype q\ [England:] near Norwich, 
Felthorpe (CM, Norwich) [examined]. 

Identity. Junior synonym of Gelis cursitans (Fabricius) (Horstmann, 1972 : 221). 

Hemiteles albomarginatus Bridgman, 1888 : 363. Holotype $, [England: Surrey,] Box 
Hill (CM, Norwich) [examined]. 

Identity. Junior synonym of Mastrus auriculatus (Thomson) (Horstmann, 1972 : 222). 

Hemiteles distinctus Bridgman, 1883 : 151. Holotype $, [England:] near Exeter (BMNH, 
on permanent loan from CMAG, Plymouth) [examined]. 

Identity. Acrolyta distincta (Bridgman) (Horstmann, 1972 : 220). 

Hemiteles incisus Bridgman, 1883 : 150. Holotype ?, [England: Surrey,] Chobham (lost). 
Identity. ? Mastrus incisus (Bridgman) (Horstmann, 1972 : 220). 

Hemiteles marginatus Bridgman, 1883 : 144. Lectotype $, [Scotland: Stirlingshire,] 
Clober (BMNH), by designation of Horstmann (1972 : 220) [examined]. 
Paralectotype q\ same data as lectotype (BMNH). 
Identity. Acrolyta rnarginata (Bridgman) (Horstmann, 1972 : 220). 

Hemiteles minutus Bridgman, 1886 : 340. Lectotype $, [England:] Worthing (CM, Nor- 
wich), by designation of Horstmann (1972 : 222) [not examined]. 
Paralectotypes. 3 6*. 3 9. same data as lectotype (CM, Norwich). 
Identity. Aclastus minutus (Bridgman) (Horstmann, 1972 : 222). 

Hemiteles mixtus Bridgman, 1883 : 148. Lectotype $, [England: Norfolk,] Brundall 
(CM, Norwich), by designation of Horstmann (1972 : 220) [examined]. 
Identity. Medophron mixtus (Bridgman) (Horstmann, 1972 : 220). 

Hemiteles nitidus Bridgman, 1889 : 416. Holotype $, [England: Dorset,] Chesil Beach 
(CM, Norwich) [examined]. 
Identity. ? Charitopes nitidus (Bridgman) (Horstmann, 1972 : 223). 

Hemiteles obscurus Bridgman, 1883 : 142. Lectotype <$, [England:] Norwich, Eaton 
(CM, Norwich), by designation of Horstmann (1972 : 219) [examined]. 
Identity. Gnypetomorpha obscura (Bridgman) (Horstmann, 1972 : 219). 

Hemiteles politus Bridgman, 1883 : 146. Lectotype $, [England:] near Exeter (CM, 
Norwich), by designation of Horstmann (1972 : 220) [examined]. 
Paralectotype $, [Great Britain] (CM, Norwich). 
Identity. Junior synonym of Eriplanus micator (Gravenhorst) (Horstmann, 1972 : 220). 

Hemiteles ruficaudatus Bridgman, 1883 : 149. Lectotype $, [England: Surrey,] near 
Shere (CM, Norwich), by designation of Horstmann (1972 : 220) [examined]. 
Identity. Bathythrix ruficaudatus (Bridgman) (Horstmann, 1972 : 220). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 313 

Hemiteles subannulatus Bridgman, 1883 : 147. Lectotype $, [England: Surrey,] near 
Shere (CM, Norwich), by designation of Horstmann (1972 : 220) [examined]. 
Paralectotypes. 4 $, same data as lectotype (BMNH). 
The lectotype has the tips of the antennae missing. 
Identity. Orthizema subannulatum (Biidgman) (Horstmann, 1972 : 220). 

Hemiteles submarginatus Bridgman, 1883 : 143. Lectotype ?, [England:] near Norwich, 
Mousehold (CM, Norwich), by designation of Horstmann (1972 : 220) [examined]. 
Paralectotypes. 4 <J, 2 ?, same data as lectotype (CM, Norwich). 
Identity. Acrolyta sabmarginata (Bridgman) (Horstmann, 1972 : 220). 

Herpestomus distinctus Bridgman, 1888 : 362. Holotype $, [England: Surrey,] Caterham 
(CM, Norwich) [examined]. 

Identity. Phaeogenes distinctus (Bridgman) (Perkins, 1953 : 116). 

Herpestomus striatus Bridgman, 1881 : 145. LECTOTYPE $, [England:] Norwich 
(CM, Norwich), here designated [examined]. 

Paralectotype $, same data as lectotype (CM, Norwich). 

Identity. Junior synonym of Oiorhinus pallipalpis Wesmael (Perkins, 1953 : 116). 

Hoplocryptus thomsoni Bridgman, 1881 : 154. Replacement name for Hoplocryptus 
elegans Thomson. 

Proposed, unnecessarily, because of the existence of another cryptine with the name 
elegans — Cryptus elegans Desvignes. Homonymy is not involved. 

Ichneumon heracliana Bridgman, 1884 : 421. LECTOTYPE $, [England: Norfolk,] 
Lynn [= Kings Lynn] (CM, Norwich), here designated [examined]. 
Paralectotypes. 6$, [Great Britain] (CM, Norwich). 
Identity. Barichneumon heracliana (Bridgman) (Perkins, 1953 : 116). 

Ichneumon pulchellatus Bridgman, 1889 : 411. LECTOTYPE ?, [Great Britain] (CM, 
Norwich), here designated [examined]. 

Paralectotypes. 2 q\ same data as lectotype (CM, Norwich). 

The lectotype and one paralectotype are mounted on a single card. The lectotype is the 
left-hand specimen. 

Identity. Platylabops pulchellatus (Bridgman) (Perkins, 1953 : 116). 

Ichneumon rufidorsatus Bridgman, 1888 : 361. Holotype $, [Scotland: Inverness-shire,] 
Aviemore (CM, Norwich) [examined]. 

Identity. Ichneumon rufidorsatus Bridgman (Perkins, 1953 : 116). 

Limneria (Angitia) aculeata Bridgman, 1889 : 428. Lectotype $, [Great Britain] (CM, 
Norwich), by designation of Horstmann (1969 : 455) [examined]. 
Identity. Diadegma aculeata (Bridgman) (Horstmann, 1969 : 455). 

Limneria (Angitia) albonotata Bridgman, 1889 : 427. Lectotype q\ [England: Surrey,] 
near Shiere [= Sherel (CM, Norwich), by designation of Horstmann (1972 : 227) [examined]. 
Paralectotype o* (CM, Norwich). 
Identity. Hyposoter albonotata (Bridgman) (Horstmann, 1969 : 419). 

Limneria (Angitia) annulipes Bridgman, 1889 : 424. Lectotype §, [Great Britain] 
(CM, Norwich), by designation of Horstmann (1972 : 227) [examined]. 

Identity. Junior synonym of Diadegma pusio (Holmgren) (Horstmann, 1969 : 434). 

Limneria barrettii Bridgman, 1881 : 158. Lectotype $, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 223) [examined]. 

Paralectotypes. 3 o\ 2 ?. same data as lectotype (CM, Norwich). 
Identity. Hyposoter barrettii (Bridgman) (Horstmann, 1972 : 223). 

Limneria brischkei Bridgman, 1882 : 153. Holotype $, [England: Cornwall,] Falmouth 
(BMNH, on permanent loan from CMAG, Plymouth) [examined]. 
Identity. Hyposoter brischkei (Bridgman) (Horstmann, 1972 : 223; teste M. G. Fitton). 



3H M. G. FITTON 

Limneria (Angitia) crassa Bridgman, 1889 : 425. Lectotype $, [Great Britain:] Abbott's 
Wood (CM, Norwich), by designation of Horstmann (1969 : 434) [examined]. 
Identity. Diadegma crassa (Bridgman) (Horstmann, 1969 : 434). 

Limneria distincta Bridgman, 1888 : 367. Holotype q\ [Great Britain:] Abbott's Wood 
(CM, Norwich) [examined]. 

Identity. Junior primary homonym of Limneria distincta Provancher. Valid name 
Sinophorus renominatus (Morley) (Horstmann, 1972 : 226). 

Limneria elishae Bridgman, 1884 : 426. Lectotype <j>, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1969 : 434) [examined]. 

Paralectotypes. 2 q", 1 $, same data as lectotype (CM, Norwich). 
Identity. Diadegma elishae (Bridgman) (Horstmann, 1969 : 434). 

Limneria (Omorga) fasciata Bridgman, 1889 : 422. Lectotype $, [England: West Sussex,] 
Steyning (CM, Norwich), by designation of Horstmann (1972 : 226) [examined]. 

Paralectotypes. 70*, 1 $. same data as lectotype (CM, Norwich); 1 $, [England: West 
Sussex,] near Littlehampton (CM, Norwich); 1 <$, [Scotland: Lewis,] Stornoway (CM, 
Norwich) . 

Identity. Junior synonym of Campoplex hadrocerus (Thomson) (Horstmann, 1972 : 226). 

Limneria fitchii Bridgman, 1881 : 157. Lectotype $, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 223) [examined]. 

Paralectotypes. 1 $ (CM, Norwich); 2 $ (BMNH). All same data as lectotype. 
Identity. Hyposoter fitchii (Bridgman) (Horstmann, 1972 : 223). 

Limneria kriechbaumeri Bridgman, 1882 : 151. Lectotype $, [Great Britain] (CM, 
Norwich), by designation of Horstmann (1972 : 224) [examined]. 
Paralectotypes. 2 £, same data as lectotype (CM, Norwich). 
Identity. Spudastica kriechbaumeri (Bridgman) (Horstmann, 1972 : 224). 

Limneria monticolana Bridgman, 1 88 1 : 159. Lectotype o*, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1970 : 81) [examined]. 
The lectotype has the mid and hind tarsi missing. 
Identity. Junior synonym of Meloboris gracilis Holmgren (Horstmann, 1972 : 224). 

Limneria reticulata Bridgman, 1884 : 430. Lectotype $, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1969 : 441) [examined]. 

Paralectotypes. 3 <$, 2 $, same data as lectotype (CM, Norwich). 

Identity. Junior synonym of Diadegma apostata (Gravenhorst) (Horstmann, 1969 : 441). 

Limneria rufa Bridgman, 1882 : 152. Lectotype o\ [Great Britain] (CM, Norwich), by 
designation of Horstmann (1972 : 224) [examined]. 

Paralectotypes. 1 <J (CM, Norwich); 1 $ (CMAG, Plymouth). Both same data as 
lectotype. 

Identity. Junior synonym of Hyposoter orbator (Gravenhorst) (Horstmann, 1972 : 224). 

Limneria rufata Bridgman, 1884 : 429. Lectotype $, [Great Britain] (CM, Norwich), 
by fixation of Morley (1915c : 201) (labelled as lectotype by Horstmann, 1972 : 225) [exam- 
ined] . 

Paralectotypes. 3 <J, 2 $, same data as lectotype (CM, Norwich). 
Identity. Diadegma rufata (Bridgman) (Horstmann, 1972 : 225). 
Limneria ruficornis Bridgman, 1884 : 429. Lectotype $, [England:] Worcestershire 
(CM, Norwich), by designation of Horstmann (1972 : 224) [examined]. 
Paralectotypes. 2 $, same data as lectotype (CM, Norwich). 

Identity. Junior synonym of Meloboris alternans (Gravenhorst) (Horstmann, 1972 : 224). 
Limneria (Angitia) scotiae Bridgman, 1889 : 426. Lectotype $, [Scotland:] Shetland 
(CM, Norwich), by designation of Horstmann (1969 : 435) [examined]. 
Paralectotypes. 5 <J, same data as lectotype (CM, Norwich). 
Identity. Diadegma scotiae (Bridgman) (Horstmann, 1969 : 435). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 315 

Limneria (Omorga) submarginata Bridgman, 1889 : 423. Lectotype 9. [England:] 
Cambridgeshire (CM, Norwich), by designation of Horstmann (1972 : 226) [examined]. 
Paralectotypes. 2 <J, 2 9, same data as lectotype (CM, Norwich). 
Identity. Junior synonym of Campoplex coracinus (Thomson) (Horstmann, 1972 : 226). 

Limneria (Anilasta) teucrii Bridgman, 1889 : 429. Lectotype 9. [Great Britain] (CM, 
Norwich), by fixation of Morley (1915c : 212) (labelled as lectotype by Horstmann, 1972 : 227) 
[examined] . 

Paralectotypes. 3 q* (CM, Norwich) ; 2 9 (CMAG, Plymouth). All same data as lectotype. 
Identity. Junior synonym of Hyposoter barrettii (Bridgman) (Horstmann, 1972 : 227). 

Limneria tripunctata Bridgman, 1886 : 351. Holotype ?, [England: London,] Peckham 
(CM, Norwich) [examined]. 

Identity. Diadegma tripunctata (Bridgman) (Horstmann, 1972 : 226). 

Limneria variabilis Bridgman, 1886 : 352. Lectotype $, [England: Cambridgeshire,] 
Wicken (CM, Norwich), by designation of Horstmann (1972 : 226) [examined]. 
Paralectotypes. 6 <J, 5 9, same data as lectotype (CM, Norwich). 
Identity. Campoplex variu "'•"., (Bridgman) (Horstmann, 1972 : 226). 

Lissonota distincta Bridgman, 1889 : 437. Holotype 9. [England: West Sussex,] Arundel 
Park (CM, Norwich) [examined]. 

Identity. Lissonota distincta Bridgman. 

Lissonota fletcheri Bridgman, 1882 : 163. LECTOTYPE 9. [England:] Worcestershire 
(CM, Norwich), here designated [examined]. 

Paralectotype $, same data as lectotype (CM, Norwich). 
Identity. Lissonota fletcheri Bridgman. 

Lissonota formosa Bridgman, 1888 : 378. Holotype 9. [Great Britain] (CM, Norwich) 
[examined] . 

Identity. Lissonota formosa Bridgman. 

Lissonota nitida Bridgman, 1886 : 371. Holotype 9. [Great Britain] (CM, Norwich) 
[examined] . 

The holotype lacks the tips of the hind tarsi. 
Identity. Lissonota nitida Bridgman. 

Lissonota obsoleta Bridgman, 1889 : 436. Syntypes 1 <J, i 9. [England: West Sussex,] 
near Littlehampton (lost). 

Identity. Unknown, the name remains a nomen dubium. 

Lissonota rufomedia Bridgman, 1886 : 370. LECTOTYPE 9, [Scotland:] Rannoch (CM, 
Norwich), here designated [examined]. 

Paralectotypes. 5 6*. 5 9, same data as lectotype (CM, Norwich); 1 9, [England: West 
Sussex,] Worthing (CM, Norwich). 

Identity. Junior synonym of Lissonota variabilis Holmgren (Aubert, 1972 : 6). 

Lissonota subaciculata Bridgman, 1886 : ^ji. Holotype 9. [Great Britain] (CM, Norwich) 
[examined] . 

Identity. Lissonota subaciculata Bridgman. 

Lissonota transversa Bridgman, 1889 : 438. Holotype 9, [England:] Norwich (CM, Norwich) 
[examined]. 

Capron's specimens are not regarded as syntypes because of the way in which Bridgman's 
statement on material is worded. 

Identity. Junior synonym of Lissonota folii Thomson (Aubert, 19696 : 90). 

Lissonota trochanterata Bridgman, 1889 : 438. Holotype 9. [England:] near Lincoln 
(CM, Norwich) [examined]. 

Identity. Junior synonym of Lissonota variabilis Holmgren (Aubert, 1972 : 6). 



316 M. G. FITTON 

Mesochorus aciculatus Bridgman, 1881 : 162. Syntypes 2 $, [Great Britain] (lost). 

Two specimens in the Bridgman collection (CM, Norwich) labelled 'Syntypes aciculatus' 
by K. Horstmann were, according to data on the mount, reared in 1882 and therefore cannot 
be types. The types are not in the Bignell collection (CMAG, Plymouth). Two males 
from the Bignell collection, where they were incorrectly labelled as types of aciculatus, are 
now in the BMNH collection (on permanent loan from CMAG, Plymouth). 

Identity. ? Mesochorus aciculatus Bridgman. 

Mesochorus facialis Bridgman, 1884 : 431. LECTOTYPE 9, [Great Britain] (CM, Nor- 
wich), here designated [examined]. 

Paralectotypes. 3 q", i $ (CM, Norwich); 8 6\ 1 $ (CMAG, Plymouth). All same data 
as lectotype. 

The lectotype is on the same mount as 1 a* and 1 $ paralectotypes. 

Identity. Mesochorus facialis Bridgman. 

Mesochorus formosus Bridgman, 1882 : 154. LECTOTYPE $, [Great Britain] (CM, 
Norwich), here designated [examined], 

Paralectotypes. 4 $ (BMNH) ; 4 $ (CMAG, Plymouth). All same data as lectotype. 
Identity. Mesochorus formosus Bridgman. 

Mesochorus hirsutus Bridgman, 1883 : 168. LECTOTYPE 6\ [England: Surrey,] Shere 
(CM, Norwich), here designated [examined]. 

? Paralectotypes. 2 $, 1 $, same data as lectotype (BMNH). 

The ? paralectotypes are from Capron's collection and are labelled 'dorsalis Holmg.'. 

Identity. Junior synonym of Asiiphromma dorsale (Holmgren) (Thomson, 1886 : 328). 

Mesochorus pectinipes Bridgman, 1883 : 166. Holotype q\ [England:] Norwich, Norkhouse 
Lane (CM, Norwich) [examined]. 

Identity. Mesochorus pectinipes Bridgman. 

Mesoleius attenuatus Bridgman, 1888 : 371. Holotype q\ [Great Britain] (CM, Norwich) 
[examined] . 

The holotype has the tips of the antennae and fore and mid tarsi missing. 
Identity. Alexetev attenuatus (Bridgman) comb. n. 

Mesoleius bignellii Bridgman, 1881 : 163. Holotype $, [Great Britain] (BMNH, on 
permanent loan from CMAG, Plymouth) [examined]. 

The holotype lacks the basal part of the first segment of the gaster. 

Identity. Perispitda bignellii (Bridgman). Generic placement established by Morley 
(191 ic : 231) here confirmed. 

Mesoleius caninae Bridgman, 1886 : 363. LECTOTYPE $, [England:] Norwich (CM, 

Norwich), here designated [examined]. 

Paralectotype o\ same data as lectotype (CM, Norwich). 

Identity. ? Otlophorus caninae (Bridgman) comb. n. (det. H. K. Townes). 

Mesoleius pini Bridgman, 1882 : 156. LECTOTYPE <j>, Scotland (CM, Norwich), here 
designated [examined]. 

Paralectotypes. 2 o\ same data as lectotype (CM, Norwich). 
The lectotype has the tips of the mid and hind tarsi missing. 

Identity. Lamachus pini (Bridgman). Generic placement established by Schmiedeknecht 
(1914 : 2905) here confirmed. 

Mesoleptus marginatus Bridgman. 1886 : 356. Holotype <J, [England:] near Norwich, 
Brundall (CM, Norwich) [examined]. 

Identity. Hadrodactylus marginatus (Bridgman) comb. n. 

Mesoleptus scutellatus Bridgman, 1886 : 356. Holotype <$ [not $ as stated by Bridgman], 
[England:] Norfolk, Horning Ferry (CM, Norwich) [examined]. 

Identity. Syn. n. of Gunomeria macrodactyla (Holmgren, 1855 : 68 (Mesoleptus)), lecto- 
type $, Sweden (NR, Stockholm) [not examined]. 



WESTERN PALAEARCTIC ICHNEUMONIDAE 317 

Monoblastus caproni Bridgman, 1882 : 159. LECTOTYPE $ [the syntypes are not all 
males as stated by Bridgman], [England: Surrey,] Shere (CM, Norwich), here designated 
[examined]. 

Paralectotypes. 2 <J, 1 $ (CM, Norwich); 3 6*. 3 ? (BMNH). All same data as lectotype. 

Morley's type restriction (1911c : 287) is not accepted because he did not label any one 
specimen as type and all of his specimens (from Capron's collection, now in the BMNH) 
are mounted on a single card. Further, since there is no definite evidence that Bridgman 
returned any of the syntypes to Capron a lectotype has been designated from the specimens 
in Norwich. 

Identity. Rhorus caproni (Bridgman) comb. n. 

Nemeritis rufipes Bridgman, 1883 : 166. Holotype $, [England: Surrey,] Shere (CM, 
Norwich) [examined]. 

Identity. Junior secondary homonym of Campoplex rufipes (Gravenhorst). Valid name 
Campoplex ruficoxa (Thomson) (Horstmann, 1972 : 224). 

Perilissus fumatus Bridgman, 1880a : 54. LECTOTYPE <$, [England:] near Norwich, 
Earlham (CM, Norwich), here designated [examined]. 
Paralectotype o\ same data as lectotype (CM, Norwich). 
Identity. Junior synonym of Labrossyta scotopterus (Gravenhorst) (Morley, 1911c : 131). 

Perilissus minutus Bridgman, 1888 :37c LECTOTYPE °, [England: Surrey,] Shere 
(CM, Norwich), here designated [examined]. 

Paralectotypes. 1 o* (CM, Norwich) ; 3 q", 2 $ (BMNH). All same data as lectotype. 
The lectotype and one male paralectotype are mounted on one card. The lectotype is 
the left hand specimen. 

Identity. Lathrolestes minutus (Bridgman) comb. n. 

Perilissus triangulatus Bridgman, 1886 : 362. LECTOTYPE $, [England:] Dulwich 
(CM, Norwich), here designated [examined]. 

Paralectotype <3\ [England:] Peckham (CM, Norwich). 

The lectotype has the tips of the antennae missing. 

Identity. Syn. n. of Azelus erythropalpus (Gmelin, 1790 : 2707 (Ichneumon)), type(s) 
[?sex] (destroyed). 

Pezomachus brevis Bridgman, 1883 : 162. Holotype?, [England:] Dover (BMNH) [exam- 
ined] . 

The holotype has the mid femora and tibiae missing and parts of other appendages have 
been eaten by beetles. This specimen was referred to as the doubtful holotype by Horstmann 
(1972 : 221) but in addition to the data given by him it bears a label (in Bridgman's hand- 
writing) 'Not in Forster ?n. sp. Have taken . . . [rest illegible, 1 or 2 words]' with 'P. brevis 
n. sp.' on the reverse. 

Identity. Gelis brevis (Bridgman) (Horstmann, 1972 : 221). 
Pezomachus costatus Bridgman, 1886 : 341. Syntypes 2 $, [England:] Norwich, Norkhouse 
Lane (CM, Norwich) [examined]; 2 $, [England: Surrey,] Shere (CM, Norwich) [examined]; 
3 $, [England: Surrey,] Shere (BMNH) [examined]. 

Identity. Junior synonym of Gelis kiesenwetteri (Foerster) (Horstmann, 1972 : 222). 
Pezomachus foersteri Bridgman, 1886 : 343. Syntypes 1 $, [England: Surrey,] Shere 
(CM, Norwich) [examined]; 3 $, same data (BMNH) [examined]. 
Identity. Gelis foersteri (Bridgman) (Horstmann, 1972 : 222). 
Pezomachus hieracii Bridgman, 1883 : 162. Syntypes 1 <j>, [England:] Liverpool (CM, 
Norwich) [examined] ; 4 $, [Great Britain] (BMNH, on permanent loan from CMAG, 
Plymouth) [examined]. 

Identity. Gelis hieracii (Bridgman) (Horstmann, 1972 : 222). 
Pezomachus vagantiformis Bridgman, 1886:342. Syntypes 1 ?, [England: Surrey,] 
Shere (CM, Norwich) [examined]; 7 <J>, same data (BMNH) [examined]. 
Identity. Gelis vagantiformis (Bridgman) (Horstmann, 1972 : 222). 



318 M. G. FITTON 

Phaeogenes formosus Bridgman, 1881 : 149. Holotype $, [England:] Norwich (CM, 
Norwich) [examined]. 

Identity. Junior synonym of Aethecerus longulus Wesmael (Perkins, 1953 : 116). 

Phaeogenes nitidus Bridgman, 1886 : 337. LECTOTYPE $, [Great Britain] (CM, Norwich), 
here designated [examined]. 

Paralectotype o*. same data as lectotype (CM, Norwich). 

Identity. Junior synonym of Cratichneumon magus (Wesmael) (Perkins, 1953 : 116). 

Phaeogenes similis Bridgman, 1881 : 148. Holotype?, [Great Britain] (lost). 

The only specimen standing under this name in the Bridgman collection (labelled 'TYPE 
= Thyraeella collaris Grav. $ det. J. F. Perkins 1951') was, according to Bridgman's notebook, 
captured on 28 May 1885 and therefore cannot be the holotype. 

Identity. Junior synonym of Thyraeella collaris (Gravenhorst) (Perkins, 1953 : 116. 
But see note above). 
Phrudus monilicornis Bridgman, 1886 : 361. LECTOTYPE $, [England: Surrey,] Shere 
(CM, Norwich), here designated [examined]. 

Since Bridgman's description suggests that he had only one male and one female before 
him a further fourteen specimens from Capron's collection (BMNH) are not regarded as 
paralectotypes. 

Identity. Phrudus monilicornis (Bridgman), combination established by Dalla Torre 
(1901 : 329). Phrudus Bridgman is a junior synonym and homonym of Phrudus Foerster. 

Phygadeuonflavopunctatus Bridgman, 1889 : 414. Lectotype $, [England:] near Norwich, 
Mousehold (CM, Norwich), by designation of Horstmann (1972 : 223) [examined]. 
Paralectotypes. 2 <J, same data as lectotype (CM, Norwich). 
Identity. Polytribax flavopunctatus (Bridgman) (Horstmann, 1972 : 223). 

Phygadeuon marshalli Bridgman, 1883 : 141. LECTOTYPE <J, [England: Devon,] 
Barnstaple [not Bugbrooke as stated by Bridgman] (BMNH), here designated [examined]. 

Paralectotype o*. same data as lectotype (BMNH). 

The two syntypes were standing in the Fitch collection under the labels 'procerus Gr. 
var. 2. n. sp.' (in Marshall's handwriting. Now attached to the lectotype.) and '389. Marsh- 
alli, Bdg.' (from Morley's catalogue) together with two circular labels marked 'S.T.'. The 
locality is indicated on the undersides of the card mounts of the specimens as 'B' which 
(Morley, 1915a) means Barnstaple. Presumably Bridgman misinterpreted this as indicating 
Bugbrooke. 

The specimen referred to by Horstmann (1972 : 219) as the ? Holotype has no type status. 

The lectotype and paralectotype are the same specimens as the lectotype and paralectotype, 
respectively, of Phygadeuon marshalli Bridgman & Fitch. 

Identity. Junior primary homonym and synonym of Phygadeuon marshalli Bridgman & 
Fitch. Valid name ITheroscopus marshalli (Bridgman & Fitch). 

Phygadeuon (Microcryptus) rufoniger Bridgman, 1889 : 415. Lectotype 9. [England: 
East Sussex,] Ashdown Forest (CM, Norwich), by designation of Horstmann (1972 : 223) 
[examined] . 

Paralectotype $, same data as lectotype (CM, Norwich). 

Identity. Junior synonym of Aptesis bifrons (Gmelin) (Horstmann, 1972 : 223). 

Phygadeuon rusticellae Bridgman, 1886 : 337. Lectotype $, [England:] Bognor (CM, 
Norwich), by designation of Horstmann (1972 : 222) [examined]. 
Paralectotypes. 4 <J, 7 ?, same data as lectotype (CM, Norwich). 
Identity. Phygadeuon rusticellae Bridgman. 

Phygadeuon tarsatus Bridgman, 1881 : 150. Lectotype $, [Scotland:] Arran, near Lamlash 
(BMNH), by designation of Townes, Momoi & Townes (1965 : 160) [examined]. 
Paralectotypes. 1 $ (BMNH); 1 $ (CM, Norwich). Both same data as lectotype. 
Identity. Aconias tarsatus (Bridgman) (Townes, Momoi & Townes, 1965 : 160). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 319 

Pimpla similis Bridgman, 1884 : 433. LECTOTYPE $, [England:] near Norwich, Brundall 
(CM, Norwich), here designated [examined]. 

Paralectotypes. 3 $, [Great Britain] (CM, Norwich). 

Identity. Junior synonym of Sea mbus nigricans (Thomson) (Oehlke, 1967 : 3). 

Platylabus transversus Bridgman, 1889 : 412. LECTOTYPE q", [Great Britain] (CM, 
Norwich), here designated [examined]. 

Paralectotype q\ same data as lectotype (CM, Norwich). 
Identity. Platylabus transversus Bridgman (Perkins, 1953 : 116). 

Polyblastus utiicinctus Bridgman, 1889 : 433. Holotype $, [England:] Devon, Slapton 
(CM, Norwich) [examined]. 

Identity. Syn. n. of Polyblastus cothurnatus (Gravenhorst, 1829b : 285 (Tryphon)), holo- 
type $, Warmbrunn (lost). 

Polysphincta subrufa Bridgman, 1888 : 377. LECTOTYPE $>, [England: Norfolk,] Lynn 
[= Kings Lynn] (CM, Norwich), here designated [examined]. 

Paralectotype $, [Scotland: Inverness-shire,] Aviemore (CM, Norwich). 
Identity. Ctenochira subrufa (Bridgman) comb. n. 

Prionopoda glaber Bridgman, 1886 : 360. LECTOTYPE $, [England:] Norwich (CM, 
Norwich), here designated [examined]. 

Paralectotypes. 4 q\ same data as lectotype (CM, Norwich). 
Identity. Rhorus glaber (Bridgman) comb. n. 

Sagaritis fasciata Bridgman, 1888 : 366. Lectotype ?, [England: Norfolk,] Horning Ferry 
(CM, Norwich), by designation of Horstmann (1972 : 226) [examined]. 
Paralectotypes. 1 5*, 1 $, same data as lectotype (CM, Norwich). 
Identity. Campoletis fasciata (Bridgman) (Horstmann, 1972 : 226). 

Sagaritis incisa Bridgman, 1883 : 165. Lectotype ?, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 224) [examined]. 
Paralectotype $, same data as lectotype (CM, Norwich). 
Identity. Campoletis incisa (Bridgman) (Horstmann, 11)72 : 224). 

Sagaritis postica Bridgman, 1886 : 350. Holotype $, [England,] near Colchester (lost). 
This specimen is presumably also the type of Sagaritis postica Bridgman & Fitch. 
Identity. Junior primary homonym and synonym of Sagaritis postica Bridgman & Fitch. 
Valid name Campoletis postica (Bridgman & Fitch) (Horstmann, 1972 : 226). 

Sagaritis punctata Bridgman, 1886 : 349. Lectotype $, [Great Britain] (CM, Norwich), 
by designation of Horstmann (1972 : 225) [examined]. 

Paralectotypes. 5 6*. 2 ?. same data as lectotype (CM, Norwich). 
Identity. Campoletis punctata (Bridgman) (Horstmann, 1972 : 225). 

Thaumatotypus billupsi Bridgman, 1882 : 145. Holotype $, [England: Surrey,] Box Hill 
[not Burford Bridge as stated by Bridgman] (BMNH) [examined]. 

The tips of the antennae are missing from the holotype. There is a slight possibility that 
this specimen is not the holotype. 

Identity. Junior synonym of Polyaulon paradoxus (Zetterstedt) (Horstmann, 1972 : 219). 

Theroscopus niger Bridgman, 1883 : 152. Holotype $, [Scotland: Inverness-shire,] Kin- 
gussie (BMNH) [examined]. 

Identity. ?Mastrus niger (Bridgman) (Horstmann, 1972 : 221). 

Thersilochus carinatus Bridgman, 1889 : 430. Holotype $, [England:] Norwich, Lucks- 
wood Lane (CM, Norwich) [examined]. 

Identity. Junior synonym of Diaparsis carinifer (Thomson) (Horstmann, 1971 : 103). 

Thersilochus marginatus Bridgman, 1886 : 354. Lectotype $, [England:] near Plymouth 
(CM, Norwich), by designation of Horstmann (1971 : 93) [examined]. 



320 M. G. FITTON 

There are four specimens under this name in the Bignell collection (CMAG, Plymouth), 
of which two may be paralectotypes. 

Identity. Probles marginatus (Bridgman) (Horstmann, 1971 : 93). 

Thersilochus minutus Bridgman, 1889 : 431. Lectotype $, [England: Surrey,] Shere 
(CM, Norwich), by designation of Horstmann (1971 : 74) [examined]. 
Paralectotype q", same data as lectotype (CM, Norwich). 
Identity. Phradis minutus (Bridgman) (Horstmann, 1971 : 74). 

Thersilochus nitidus Bridgman, 1889 : 430. PHolotype $, [England: Wiltshire,] Nunton 
(BMNH) [examined]. 

The Pholotype was labelled as such by Horstmann. It also bears labels: 'N' (on underside 
of mount), 'Wilts: Nunton, nr. Salisbury. Marshall Coll. B.M. 1904-120.' (printed), and 
'B.M. Coll. 1949 under Thersiloch. nitidus, Bdgm.'. 

There is no direct evidence that this is the holotype. 

Identity. Pygmaeolus nitidus (Bridgman). Type-species of Pygmaeolus Hellen. 

Thymaris fasciatus Bridgman, 1886 : 348. Holotype $, [England:] Norfolk (CM, Norwich) 
[examined] . 

This specimen is also the holotype of Thymaris fasciatus Bridgman & Fitch. 

Identity. Junior primary homonym and synonym of Thymaris fasciatus Bridgman & 
Fitch. Valid name Cymodusa fasciatus (Bridgman & Fitch). 



Bridgman, J. B. & Fitch, E. A. 

Bridgman & Fitch published a series of keys to British ichneumonids which 
include a number of species described as new by Bridgman. Unfortunately in 
the following cases the keys appeared before Bridgman's formal descriptions and 
therefore validate the names concerned. 

Anomalon cylindricum Bridgman & Fitch, 1884 (October) : 224. LECTOTYPE 9, [Great 
Britain] (PEM, London), here designated [examined]. 

Paralectotype $, same data as lectotype (PEM, London). 

The lectotype and paralectotype are the same specimens as the lectotype and paralectotype, 
respectively, of Anomalon cylindricum. Bridgman. 

Identity. Barylypa cylindricum (Bridgman & Fitch) comb. n. 

Anomalon minutum Bridgman & Fitch, 1884 (October) : 224. LECTOTYPE $, [England: 
Hampshire,] New Forest (CM, Norwich), here designated [examined]. 

Paralectotypes. 2 <3\ 5 $. same data as lectotype (CM, Norwich). 

The lectotype and paralectotypes are the same specimens as the lectotype and paralectotypes, 
respectively, of Anomalon minutum Bridgman. 

Identity. Agrypon minutum (Bridgman & Fitch) (det. I. D. Gauld). 

Campoplex costulatus Bridgman & Fitch, 1885a : 17. Type(s) a*, [Great Britain] (lost). 
The type-material of this species is presumably the same as that of Campoplex costulatus 
Bridgman. 

Identity. Junior synonym of Dusona anceps (Holmgren). Synonymy of Bridgman's 
species established by Hinz (1972 : 47). 

Campoplex femorator Bridgman & Fitch, 1885a : 17. LECTOTYPE $, [Great Britain] 
(CM, Norwich), here designated [examined]. 

Paralectotypes. 1 $ (CM, Norwich) ; 2 g, 2 $ (BMNH). All same data as lectotype. 

The lectotype and paralectotypes are the same specimens as the lectotype and paralecto- 
types, respectively, of Campoplex femorator Bridgman. 

Identity. Junior synonym of Dusona polita (Foerster). Synonymy of Bridgman's species 
established by Hinz (1972 : 47). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 321 

Campoplex punctatus Bridgman & Fitch, 1885a : 16. Holotype 6\ [Great Britain] (CM, 
Norwich) [examined]. 

This specimen is also the holotype of Campoplex punctatus Bridgman. 

Identity. Junior synonym of Dusona petiolator (Fabricius). Synonymy of Bridgman's 
species established by Hinz (1972 : 47). 

Phygadeuon marshalli Bridgman & Fitch, 1882a : 228. LECTOTYPE <$, [England: 
Devon,] Barnstaple (BMNH), here designated [examined]. 

Paralectotype $, same data as lectotype (BMNH). 

Details of the type-material were given by the authors in the next paper in the series 
(18826 : 275). The lectotype and paralectotype are the same specimens as the lectotype 
and paralectotype, respectively, of Phygadeuon marshalli Bridgman. 

Identity. ITheroscopus marshalli (Bridgman & Fitch) comb. n. 

Sagaritis postica Bridgman & Fitch, 18856 : 101. Type(s) [?sex], [Great Britain] (lost). 

The type-material of this species is presumably the same as that of Sagaritis postica 
Bridgman. 

Identity. Campoletis postica (Bridgman & Fitch). Generic placement established by 
Horstmann (1972 : 226) for Bridgman's species. 

Thymaris fasciatus Bridgman & Fitch, 18856 : 100. Holotype $, [England:] Norfolk 
(CM, Norwich) [examined]. 

This specimen is also the holotype of Thymaris fasciatus Bridgman. 
Identity. Cymodusa fasciatus (Bridgman & Fitch) comb. n. 



Cameron, P. 

Cameron described a large number of Ichneumonidae but only one species from 
the western Palaearctic. 

Crypturus fulvipes Cameron, 1901 : 330. Holotype $, Gibraltar (BMNH) [examined]. 
The holotype has lost the tips of the hind tarsi. 
Identity. Latibulus fulvipes (Cameron) comb. n. 



Capron, E. 

Pezomachus pilosus Capron, 1888 : 217. LECTOTYPE $, [England: Surrey,] Shiere 
[=Shere] (BMNH), here designated [examined]. 

Paralectotypes. 2 $, same data as lectotype (BMNH) . 

The three syntypes were mounted on a single card with 'pilosus Capron' (in Capron's 
handwriting) on a strip of paper glued to the card. The paralectotypes remain on this card. 
The lectotype has been remounted (? by J. F. Perkins) on a card point. 

Identity. Gelis pilosus (Capron). Generic placement established by Ceballos (1925 : 19°) 
here confirmed. 
Pimpla varicauda Capron, 1888 : 217. LECTOTYPE $, [England : Surrey,] Shiere [ = Shere] 
(BMNH), here designated [examined]. 

Paralectotype $, same data as lectotype (BMNH). 

The lectotype and paralectotype are mounted on a single card with 'varicauda. Capron' 
(in Capron's handwriting) and 'EMM 1888, p. 218, $.' (in Morley's handwriting) on a strip 
of paper glued to the card. The lectotype is the left-hand specimen. 

Identity. Junior synonym of Stilbops vetula (Gravenhorst) . Synonymy indicated by 
Morley (1908 : 173). Generic placement here confirmed. 



322 M. G. FITTON 

Curtis, J. 

Neboiss (1963) gives some notes on the Curtis collection and associated manuscript 
material. 

Curtis attributes a number of names proposed in his British Entomology to other 
authors (notably Haliday) but I consider Curtis to be responsible for the descriptions 
as published and therefore as author of the species concerned. 

Agriotypus armatus Curtis, 1832 : 389. LECTOTYPE $, [Scotland:] near Lanark, on 
the [River] Clyde (BMNH), here designated [examined]. 

Paralectotypes. 2 $, 4 <j>, same data as lectotype (NMV, Melbourne). 
Identity. Agriotypus armatus Curtis. 

Alomya victor Curtis, 1826 : 120. LECTOTYPE $, [Scotland: Perthshire,] Dunkeld 
(NMV, Melbourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 
The lectotype lacks the tips of the hind tarsi and the gaster is detached. 
Eleven further specimens [not examined] in the Curtis collection may be paralectotypes. 
Identity. Junior synonym of Alomya debellator (Fabricius). 

Anomalon vesparum Curtis, 1828 : 198. LECTOTYPE $, [Great Britain] (NMV, Mel- 
bourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 

The lectotype lacks the mid legs, except the left coxa, and the tips of the hind tarsi. 
Five further specimens [not examined] in the Curtis collection may be paralectotypes. 
Identity. Spliecophaga vesparum (Curtis). Type-species of Sphecophaga Westwood. 

Banchus far rani Curtis, 1836 : 588. LECTOTYPE $, [Ireland:] Connemara (NMV, Mel- 
bourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 
The lectotype lacks the tips of the antennae. 

A further specimen [not examined] in the Curtis collection is probably the paralectotype. 
Identity. Junior synonym of Banchus volutatorius (Linnaeus) (det. M. G. Fitton). 

Bassus athaliaeperda Curtis, i860 : 53. Holotype q\ [Great Britain] (NMV, Melbourne) 
[examined] . 

The holotype lacks the tips of the hind tarsi. 
Identity. ? Arbelus athaliaeperda (Curtis) comb. n. 

Cremastus buoliana Curtis, 1854 : 60. LECTOTYPE $, [England:] Clifton Nursery 
(NMV, Melbourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 
Identity. Temelucha buoliana (Curtis) comb. n. 

Cryptus bellosus Curtis, 1837 : 668. LECTOTYPE $, [Great Britain] (NMV, Melbourne), 
here designated (labelled as 'type' by J. F. Perkins) [examined]. 
The lectotype lacks the tips of the hind tarsi. 

A further specimen [not examined] in the Curtis collection is probably the paralectotype. 
Identity. Junior synonym of Aritranis signatorius (Fabricius) (Morley, 1907 : 294). 

Euceros albitarsus Curtis, 1837 : 660. Holotype <J, [England:] Northamptonshire, near 
Milton (NMV, Melbourne) [examined]. 

The holotype lacks the apices of the antennae and all tarsi; and the gaster except the 
first two segments. 

Identity. Euceros albitarsus Curtis. 

Ichneumon atropos Curtis, 1828:234. LECTOTYPE $, [England:] ?Kent, Rochester 
or Darent[h] Wood (BMNH), here designated (labelled by C. Morley) [examined]. 

Although there is now little evidence to support Morley's assumption (1902 : 118) that 
the BMNH specimen is an original syntype, it is here designated as lectotype. Curtis's 
statement 'In the Cabinets of Mr. Davis and Mr. S. Sulivan.' is taken to exclude any of his 
own material from the type-series. There are 2 $ in the Curtis collection, conspecific with 
the lectotype. 



WESTERN PALAEARCTIC ICHNEUMONIDAE 323 

Identity. Junior synonym of Callajoppa exaltatoria (Panzer) (Morley, 1902 : 118, as 



r t US). 

Lampronota crenicornis Curtis, 1832 : 407. LECTOTYPE $, Ireland (NMI, Dublin), 
here designated [examined]. 

Paralectotype q\ same data as lectotype (NMI, Dublin). 

The lectotype and paralectotype are from the Haliday collection. They were among 
material identified as crenicornis by Morley (19136 : 263) who attributed the species to 
Haliday, 1838. Therefore Morley's 'type' has no status. 

Identity. Junior synonym of Cylloceria caligata (Gravenhorst) (Morley, 1908 : 255, as 
Lampronota) . 

Mesochorus basalts Curtis, 1833 : 464. LECTOTYPE $, [England: Hampshire,] New 
Forest (NMV, Melbourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 
The lectotype lacks the tips of the antennae. 
Identity. Mesochorus basalis Curtis. 

Mesochorus fulgurans Curtis, 1833 : 464. Holotype $, Ireland (NMV, Melbourne) [exam- 
ined] . 

Identity. Mesochorus fulgurans Curtis. 

Mesochorus olerutn Curtis, 1833 : 464. LECTOTYPE ?, [Great Britain] (NMV, Mel- 
bourne), here designated [examined]. 

Paralectotype q\ same data as lectotype (NMV, Melbourne) . 
Identity. Mesochorus olerum Curtis. 

Mesochorus sericans Curtis, 1833 : 464. LECTOTYPE 9, Ireland: ?near Belfast (NMI, 
Dublin), here designated [examined]. 

Paralectotype <$, same data as lectotype (NMI, Dublin). 

The lectotype and paralectotype were standing under Mesochorus in the Haliday collection. 
There can be no doubt that they are syntypes of Curtis's species. 

Identity. Astiphromma sericans (Curtis) comb. n. 

Mesochorus splenium Curtis, 1833 : 464. Holotype <$, [England: Hampshire,] New Forest 
(NMV, Melbourne) [examined]. 

The holotype is badly damaged: the head, gaster and appendages are detached from the 
thorax. 

Identity. Astiphromma splenium (Curtis) comb. n. 

Mesochorus sylvarum Curtis, 1833 : 464. LECTOTYPE $, [Great Britain] (NMV, 
Melbourne), here designated [examined]. 

? Paralectotype <£, same data as lectotype (NMV, Melbourne). 

The Pparalectotype does not agree entirely with the original description, notably in the 
colour of the mesoscutum. 

Identity. Mesochorus sylvarum Curtis. 

Mesoleptus gracilipes Curtis, 1837 : 644. Holotype $ [not 3* as stated by Curtis], [Scotland:] 
near Lanark (NMV, Melbourne) [examined]. 
Identity. Alexeter gracilipes (Curtis) comb. n. 

Mesoleptus speciosus Curtis, 1837 : 644. Holotype $, [England:] Coomb-wood (lost). 
Identity. Unknown, the name remains a nomen dubium. 

Mesoleptus ventralis Curtis, 1837 : 644. LECTOTYPE q\ [Great Britain] (NMV, Mel- 
bourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 

A further three specimens [not examined] in the Curtis collection may be paralectotypes. 
Identity. Hadrodactylus ventralis (Curtis) comb. n. 

Mesoleptus waltoni Curtis, 1837 : 644. LECTOTYPE q\ [England:] Yorkshire, Knares- 
borough, by the River Nidd (NMV, Melbourne), here designated (labelled as 'type' by J. F. 
Perkins) [examined]. 



324 M. G. FITTON 

The lectotype lacks antennae, most of the forelegs and most of the hind tarsi. 

A further specimen [not examined] in the Curtis collection may be a paralectotype. 

Identity. Junior synonym of Sympherta fuscicornis (Gmelin) (Morley, 1911c : 237, as 

Catoglyptus) . 

Peltastes pini Curtis, 1824 : 4. Syntypes q* $, [England:] Hampshire, near Ringwood 

(lost). 

Although there are three specimens (1 $, 2 $ Metopius dentatus (Fabricius)) standing under 
dentatus/pini in the Curtis collection, Curtis's statement 'In the cabinet of Mr Bentley' is 
taken to exclude any of his own material from the type-series. It seems unlikely that Curtis 
received any of the original type material (Curtis, 1829 : 4). Bentley's collection has not 
been located. 

Identity. Junior synonym of Metopius dentatus (Fabricius) (Morley, 1911c : 3). 

Pimpla aethiops Curtis, 1828 : 214. LECTOTYPE ?, [Great Britain] (NMV, Melbourne), 
here designated (labelled as 'type' by J. F. Perkins) [examined]. 

A further specimen [not examined] in the Curtis collection may be a paralectotype. 
Identity. Pimpla aethiops Curtis. 

Pimpla cossivora Curtis, 1828 : 214. Type(s) [??], [Great Britain] (lost). 
Identity. Unknown, the name remains a nomen dubium. 

Stilpnus dryadum Curtis, 1832 : 388. LECTOTYPE $, Ireland: Galway (NMI, Dublin), 
here designated [examined]. 

Paralectotypes 1 ^ 1 $, same data as lectotype (NMI, Dublin). 

The lectotype and paralectotypes are from the Haliday collection. The lectotype is in 
very good agreement with Curtis's description and illustration and bears a handwritten 
label 'dryadum'. 

The lectotype lacks the tips of the antennae. 

Identity. Stilpnus dryadum Curtis. 

Therion gracilipes Curtis, 1839 : 736. Holotype o\ [Great Britain] (NMV, Melbourne) 
[examined] . 

The holotype lacks the tips of the hind tarsi. 

Identity. Agrypon gracilipes (Curtis) comb. n. (det. I. D. Gauld). 

Xylonomus gravenhorstii Curtis, 1831 : 353. LECTOTYPE $, [England:] near London 
(NMV, Melbourne), here designated (labelled as 'type' by J. F. Perkins) [examined]. 
Identity. Xorides gravenhorstii (Curtis). 



Desvignes, T. 

There has been some confusion as to the original combinations of the species 
Desvignes described in 1856. However, in the preface to this work Desvignes 
clearly states that 'The species are arranged according to the method, used in 
Professor I. L. C. Gravenhorst's Ichneumonologia Europaea, . . .' and therefore 
Desvignes' 'subgenera' are here treated as genera as Gravenhorst's have been. 

Anomalon capitatum Desvignes, 1856 : 104. LECTOTYPE q*. [England:] Yorkshire 
(BMNH), here designated [examined]. 

The lectotype lacks the tips of the hind tarsi. Although 'types' are mentioned by Morley 
(1915c : 228) only the lectotype can be found in the BMNH collection. 

Identity. Heteropelma capitatum (Desvignes) (det. I. D. Gauld). 

Anomalon interruptum Desvignes, 1856 : 106. Type(s) [?sex], [Great Britain] (lost). 
Identity. Agrypon interruptum (Desvignes) (det. I. D. Gauld). 



WESTERN PALAEARCTIC ICHNEUMONIDAE 325 

Anomalon mirabile Desvignes, 1856 : 105. Holotype 6*, [Great Britain] (BMNH) [exam- 
ined] . 

Identity. Junior synonym of Habronyx heros (Wesmael) (det. I. D. Gauld). 

Bassus albicinctus Desvignes, 1862a : 218. Holotype <J, [Great Britain] (BMNH) [exam- 
ined] . 

Identity. Junior synonym of Syrphophilus tricinctorius (Thunberg) (Dasch, 1964 : 68). 

Bassus compressus Desvignes, 1856 : 91. LECTOTYPE 9. [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotypes. 2 ?, same data as lectotype (BMNH). 
Identity. Phthorima compressa (Desvignes) (Dasch, 1964 : 218). 

Bassus flavus Desvignes, 1862a : 219. Holotype o\ [Great Britain] (BMNH) [examined]. 
Identity. Junior synonym of Homotropus tarsatorius (Panzer) (Dasch, 1964 : 140). 

Bassus frenat or Desvignes, 1862a : 218. Holotype q\ [Great Britain] (BMNH) [examined]. 
The head and the tips of the mid tarsi are missing from the holotype. 

Identity. Junior synonym of Enizemnm ornatum (Gravenhorst) (Morley, 1911c : 99, 
as Homocidus). 

Bassus maculatus Desvignes, 1862a : 216. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Identity. Junior synonym of Sussaba dorsalis (Holmgren) (Dasch, 1964 : 257). 

Bassus picitans Desvignes, 1862a : 217. LECTOTYPE £, [Great Britain] (BMNH), here 
designated [examined]. 

Identity. Junior synonym of Homotropus nigritarsus (Gravenhorst). Not a synonym 
of Tymmophorus graculus (Gravenhorst) as stated by Dasch (1964 : 78). 
Bassus planus Desvignes, 1862a : 220. LECTOTYPE ?q\ [Great Britain] (BMNH), 
here designated [examined]. 

The lectotype has the gaster and hind legs missing. 

Identity. ? Junior synonym of Homotropus dimidiatns (Schrank) (Morley, 1911c : 102). 
Bassus pulchellus Desvignes, 1862a : 221. LECTOTYPE <J. [Great Britain] (BMNH), 
here designated [examined]. 

The lectotype has the tips of the antennae and of the fore and hind tarsi missing. 
Identity. Junior primary homonym of Bassus pulchellus Holmgren. Replacement 
name Bassus desvignesii Marshall. Junior synonym of Homotropus tarsatorius (Panzer) 
(Dasch, 1964 : 140). 
Bassus rufocinctus Desvignes, 1862a : 215. LECTOTYPE <$, [Great Britain] (BMNH), 
here designated [examined]. 

The lectotype lacks the tips of the antennae. 

Identity. Junior synonym of Tymmophorus graculus (Gravenhorst) (Dasch, 1964 : 77). 
Bassus scabrosus Desvignes, 1862a : 217. Holotype o*. [Great Britain] (BMNH) [examined]. 
The holotype has lost most of its antennae and hind tarsi. 

Identity. Junior synonym of Homotropus sundevalli (Holmgren) (Morley, 1911c : 101, 
as Homocidus). 
Bassus thoracicus Desvignes, 1862a : 219. Type(s) [?sex], [Great Britain] (lost). 

Identity. Unknown, the name remains a nomen dubium. Placed by Morley (1911c : 104) 
as a synonym of Homotropus pictus (Gravenhorst). 
Campoplex henaultii Desvignes, 1856 : 98. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

The lectotype has the tip of the ovipositor broken off. 
Identity. Hyposoter henaultii (Desvignes) comb. n. 
Campoplex myrtillus Desvignes, 1856 : 99. Holotype 6", [Great Britain] (BMNH) [exam- 
ined] . 

Identity. Dusona myrtillus (Desvignes) (Hinz, 1957 : 86). 



326 M. G. FITTON 

Campoplex placidus Desvignes, 1856 : 97. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Identity. Hyposoter placidus (Desvignes) comb. n. 

Campoplex tomentosus Desvignes, 1856 : 100. Holotype $, [Great Britain] (BMNH) 
[examined] . 

Identity. Ctenopelma tomentosus (Desvignes) comb. n. 

Cryptus elegans Desvignes, 1856 : 57. LECTOTYPE <$, [Great Britain] (BMNH), here 
designated [examined]. 

The lectotype lacks the tips of the antennae and hind tarsi. 

The material in the Curtis collection cannot be found (A. Neboiss, personal communication). 

Identity. Junior synonym of Thrybius leucopygus (Gravenhorst) . 

Cryptus hopei Desvignes, 1856 : 58. Lectotype [2 sex], [? Great Britain] (lost), by fixation 
of Morley (1907 : 57). 

Paralectotypes. 2 $, [England:] Netley (?lost). 

The type-series of this species is considered to consist of the two females referred to by 
Gravenhorst (1829a : 705) (which cannot now be found in the Hope Department collections 
(UM, Oxford) but may be in the Gravenhorst collection in Wroclaw. They were almost 
certainly never in the BMNH collections as stated by Morley (1907 : 57)) together with a 
specimen (the lectotype) or specimens indicated by Desvignes as being in the BMNH collec- 
tions. The lectotype cannot now be found in the BMNH collections. 

Identity. lAptesis hopei (Desvignes) comb. n. See Acanthocryptus hopei Morley. 

Cryptus ruficeps Desvignes, 1856 : 55. Type(s) [?] $, [Great Britain] (lost). 

Morley (1907 : 292) states 'Only a pin-hole adorns the proper position of this "species" 
in the British Museum, and the type appears to be lost.'. 
Identity. Unknown, the name remains a nomen dubium. 

Cryptus rufoniger Desvignes, 1856 : 58. Holotype $, [Great Britain] (BMNH) [examined]. 
Most of the antennae, mid legs and ovipositor sheaths are missing from the holotype. 
Identity. Aritranis rufoniger (Desvignes). 

Cryptus sanguinator Desvignes, 1856 : 58. Syntypes [?] $, [Great Britain] (lost). 

Identity. Unknown, the name remains a nomen dubium. Placed by Morley (1903 : 82) 
as a synonym of Barichneumon sanguinator (Rossi). 

Cryptus tumidus Desvignes, 1856 : 56. Holotype $, [Great Britain] (BMNH) [examined]. 
The tips of the fore and mid tarsi are missing from the holotype. 

Identity. Junior synonym of Xylophritrus dispar (Thunberg) (Morley, 1907 : 326, as a 
synonym of lancifer Gravenhorst). 

Ephialtes albicinctus Desvignes, 18626 : 226. Holotype q, [? Great Britain] (BMNH) 
[examined] . 

Identity. Junior primary homonym of Ephialtes albicinctus Gravenhorst. Valid name 
Delomerista desvignesii (Marshall) (Oehlke, 1967 : 33). 

Ephialtes facialis Desvignes, 18626:226. LECTOTYPE 6*. [? Great Britain] (BMNH), 
here designated [examined]. 

Identity. Junior synonym of Lissonota catenator (Panzer) (Perkins, 1943a : 260). 

Ephialtes ruficollis Desvignes, 1856 : 88. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Identity. Liotryphon ruficollis (Desvignes) (Oehlke, 1967 : 9). 

Exetastes facialis Desvignes, 1856 : 95. Holotype 6", [Great Britain] (BMNH) [examined]. 
Identity. Junior synonym of Exetastes maurus Desvignes (Aubert, 1971 : 86). 

Exetastes femorator Desvignes, 1856 : 94. Lectotype $, [Great Britain] (BMNH), by 
designation of Townes, Momoi & Townes (1965 : 228) [examined]. 
Identity. Exetastes femorator Desvignes. 



WESTERN PALAEARCTIC ICHNEUMONIDAE 327 

Exetastes maurus Desvignes, 1856 : 95. LECTOTYPE $>, [Great Britain] (BMNH) 
here designated [examined]. 

Paralectotype $, same data as lectotype (BMNH). 

The tips of the antennae and fore tarsi are missing from the lectotype. 

Identity. Exetastes maurus Desvignes. 

Exochus globulipes Desvignes, 1856 : 45. Lectotype $, [Great Britain] (BMNH), by 
designation of Townes, Momoi & Townes (1965 : 353) [examined]. 
Townes' label is now missing from the lectotype. 
Identity. Triclistus globulipes (Desvignes) (Aeschlimann, 1973 : 240). 

Glypta bicornis Desvignes, 1856 : 74. Holotype o\ [Great Britain] (BMNH) [examined]. 

The holotype lacks the tips of the hind tarsi. 

Morley (1908 : 145) mentions 'two co-types' of this species. Unfortunately the specimen 
labelled by Morley as 'type' does not agree in coloration (the fourth segment of the gaster 
is largely red) with the original description and it is almost certainly not a type-specimen. 
The remaining specimen agrees with Desvignes description and is regarded as holotype of 
his species. 

Identity. Junior primary homonym and synonym of Glypta bicornis Boie (Morley, 1908 : 
M5)- 
Glypta fcmorator Desvignes, 1856 : 73. Holotype q\ [Great Britain] (BMNH) [examined]. 

The holotype has lost the tips of the hind tarsi. 

Identity. Glypta femorator Desvignes. 

Glypta flavipes Desvignes, 1856:75. LECTOTYPE 6", [Great Britain] (BMNH), here 
designated [examined]. 

Paralectotypes. 2$, same data as lectotype (BMNH). 

Identity. Junior synonym of Glypta nigrina Desvignes (Aubert, 1971 : 84). 

Glypta lineata Desvignes, 1856 : 76. LECTOTYPE $, [Great Britain] (BMNH), here 
designated [examined]. 

Paralectotypes. 1 <J, 1 J, same data as lectotype (BMNH). 
The lectotype lacks the tips of the antennae. 
Identity. Glypta lineata Desvignes. 

Glypta nigrina Desvignes, 1856 : 74. LECTOTYPE ?, [Great Britain] (BMNH), here 
designated [examined] . 

Identity. Glypta nigrina Desvignes. 

Glypta pedata Desvignes, 1856 : 74. LECTOTYPE $, [Great Britain] (BMNH), here 
designated [examined]. 

The tips of the fore and hind tarsi are missing from the lectotype. 
Identity. Glypta pedata Desvignes. 

Glypta ruficeps Desvignes, 1856 : 76. Holotype <j>, [Great Britain] (BMNH) [examined]. 
Identity. Junior synonym of Glypta nigrina Desvignes (Aubert, 1971 : 84). 

Hemiteles formosus Desvignes, in Smith & Desvignes, i860 : 211. LECTOTYPE $, [Great 
Britain] (BMNH), here designated [examined]. 

Identity. Junior synonym of Bathylhrix fragilis (Gravenhorst) (synonymy indicated by 
Morley, 1907 : 106). 

Hoplismenus semirufus Desvignes, 1856 : 50. Type(s) $, [Great Britain] (lost). 

The specimen (?holotype) examined by Morley (1902 : 123; 1903 : 229) and Heinrich 
(1937 : 2 79) is missing from the BMNH collection (Perkins, 1953 : 115) although a pin with 
the following labels is present 'Type', 'B.M. TYPE HYM. 3^1584', '7 HOPLISMENUS 
SEMIRUFUS', 'Hoplismenus semirufus Desv. Cat. 1856. 50. (type). = PPlatylabus Stalii 
Holmgr. Ichn. Suec. CM i.02.', 'semirufus', 'Dsvgns. 68-52'. 

Identity. Junior synonym of Platylabops apricus (Gravenhorst) (Heinrich, 1937 : 2 79'> 
Perkins, 1953 : 115). 



328 M. G. FITTON 

Ichneumon ancipiterus Desvignes, 1856 : 19. Lectotype o*. [Great Britain] (BMNH), 
by fixation of Perkins (1953 : 114) [examined]. 

The lectotype has the tips of the antennae, the forelegs and the tips of the hind tarsi missing. 
Identity. Junior synonym of Amblyteles palliatorius (Gravenhorst) (Perkins, 1953 : 114). 

Ichneumon binotatus Desvignes, 1856 : 23. Holotype o", [Great Britain] (BMNH) [exam- 
ined]. 

Identity. Junior primary homonym of Ichneumon binotatus Stephens. Junior synonym 
of Cratichneumon coruscator (Linnaeus) (Perkins, 1953 : 114). 

Ichneumon cambriensis Desvignes, 1867 : 130. LECTOTYPE $, [Wales: Dyfed,] Gellys- 
wick (BMNH), here designated [examined]. 

Paralectotype $, same data as lectotype (BMNH) . 

Perkins' (1953 : 115) type fixation is not accepted since there are two males each labelled 
'Type Hym. 3b. 1602' in the BMNH collection. The undamaged specimen is here designated 
lectotype. The paralectotype has the apex of the gaster missing. 

Identity. Junior synonym of Phaeogenes stipator Wesmael (Perkins, 1953 : 115). 

Ichneumon crassorius Desvignes, 1856 : 10. Lectotype $, [Great Britain] (BMNH), 
by fixation of Perkins (1953 : 113) [examined]. 
Paralectotype $, same data as lectotype (BMNH). 

It has not been possible to identify other members of Desvignes' type-series. 
Identity. Junior synonym of Ichneumon didymus Gravenhorst (Perkins, 1953 : 113). 

Ichneumon cubicularis Desvignes, 1856 : 15. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotype o", same data as lectotype (BMNH). 

Perkins' type fixation (1953 : 114) is not accepted since BMNH 'Type Hym. 3b 1571' 
is male and not female as stated. The female syntype has no BMNH serial number [It 
is now numbered 3B.2152.] but is probably the specimen referred to by Perkins. Since 
this is not clear the female specimen is here formally designated as lectotype. 

Identity. Junior synonym of Spilothyrateles fabvicii (Schrank) (Perkins, 1953 : 114; 
Heinrich, 1967 : 600). 

Ichneumon dubitatus Desvignes, 1856 : 20. Holotype $, [Great Britain] (BMNH) [exam- 
ined] . 

The holotype lacks the antennae, forelegs and the tips of the mid and hind tarsi. 

Identity. Junior synonym of Amblyteles palliatorius (Gravenhorst) (Perkins, 1953 : 114). 

Ichneumon flavocinctus Desvignes, 1856 : 22. Holotype 0*. [Great Britain] (BMNH) 
[examined] . 

Although Desvignes gives a range of length this is assumed to include the specimen (s) 
differentiated as Var. 1., which are specifically excluded from the syntype series under the 
International Code of Zoological Nomenclature (1961 : Article 72(b)). Therefore the only 
known syntype is regarded as holotype. 

Identity. Junior synonym of Ctenichneumon panzeri (Wesmael) (Perkins, 1953 : 114)- 

Ichneumon lautatorius Desvignes, 1856 : 18. Lectotype $, [Great Britain] (NMV, Mel- 
bourne) by fixation of Perkins (1953 : 114) [examined]. 

Paralectotype o", same data as lectotype (NMV, Melbourne). 
Identity. Ichneumon lautatorius Desvignes (Perkins, 1953 : 114)- 

Ichneumon maculiventris Desvignes, 1856 : 5. Holotype §, [England: Hampshire, 
New Forest] (BMNH) [examined]. 

The tips of the antennae are missing from the holotype. The specimen is from Stephens' 
collection where it stood as Ichneumon fasciatus Gmelin (Stephens, 1835 : 129). Both 
Morley (1903 : 170) and Perkins (1953 : 107) mistakenly attributed the name fasciatus 
to Stephens and regarded the holotype of Desvignes' species as its 'type'. 

Identity. Junior synonym of Hepiopelmus leucostigmus (Gravenhorst) (Perkins, 1953 : 113)- 



WESTERN PALAEARCTIC ICHNEUMONIDAE 329 

Ichneumon minutorius Desvignes, 1856 : 22. Lectotype $, [Great Britain] (BMNH), 
by fixation of Perkins (1953 : 114) [examined]. 

Paralectotypes. 7 <$, same data as lectotype (BMNH). 

Since variants are excluded from the syntype series under the Code (1961 : Article 72(b)) 
it has been necessary to determine which of the twenty specimens mentioned by Desvignes 
belong to his varieties 1 and 2. Sixteen specimens have been traced in the BMNH collection, 
of which eight (lectotype and paralectotypes) belong to the type form and seven and one, 
respectively, to varieties 1 and 2. 

Identity. Ichneumon minutorius Desvignes (Perkins, 1953 : 114). 

Ichneumon niveatus Desvignes, 1856 : 25. Lectotype q", [Great Britain] (BMNH), by 
fixation of Perkins (1953 : 114) [examined]. 

Paralectotypes. I2<J, same data as lectotype (BMNH). 

The lectotype has the antennae and most of the hind tarsi missing. 

Identity. Junior synonym of Oresbius arridens (Gravenhorst) comb. n. (Perkins, 1953 : 
114; Townes & Gupta, 1962 : 131; Townes, 1970a : 131). 

Ichneumon obator Desvignes, 1856 : 5. Lectotypec?, [Great Britain] (BMNH), by fixation 
of Perkins (1953 : 113) [examined]. 

There are four further males (1 ex coll. J. F. Stephens, 3 ex coll. Desvignes) in the BMNH 
collection which may be paralectotypes of this species. 

Identity. Platylabus obator (Desvignes) (Perkins, 1953 : 113). 

Ichneumon paludator Desvignes, 1854 : 44. Lectotype $, [England: London,] Hammer- 
smith Marshes (BMNH), by fixation of Perkins (1953 : 115) [examined]. 
Identity. Chasmias paludator (Desvignes) (Perkins, 1953 : 115). 

Ichneumon re lucens Desvignes, 1856 : 11. Holotype?, [Great Britain] (BMNH) [examined]. 
Identity. Junior synonym of Diphyus indocilis (Wesmael) (Perkins, 1953 : 113; Townes, 
Momoi & Townes, 1965 : 490). 

Ichneumon rubedinis Desvignes, 1856 : 29. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotypes. 2 $, same data as lectotype (BMNH). 

Since there are three specimens labelled with the serial number 'BM Cotype HYM 3b. 1569' 
it is not possible to ascertain which one Perkins (1953 : 115) intended to fix as type. A 
lectotype is therefore designated here. 

Identity. Junior synonym of Ichneumon walkeri Wesmael (Perkins, 1953 : 115)- 

Lampronota frontalis Desvignes, 1856 : 78. LECTOTYPE 6", [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotypes. 3 <$, same data as lectotype (BMNH). 

Of the four syntypes (Morley, 1908 : 215) the specimen previously bearing a BMNH 
'type' label was not selected as lectotype because of its poor condition. 

Identity. Lissonota frontalis (Desvignes). 

Lampronota fulvipes Desvignes, 1856 : 78. LECTOTYPE ?, [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotypes. 3 <J, 2 $, same data as lectotype (BMNH). 
Identity. Lissonota fulvipes (Desvignes). 

Lampronota notabilis Desvignes, 1856 : 79. Holotype <3\ [Great Britain] (BMNH) 
[examined] . 

The holotype is in poor condition and lacks most of the antennae, fore tarsi, the tips of 
the mid tarsi and all of the hindlegs except the left coxa. 
Identity. Leptacoenites notabilis (Desvignes) comb. n. 

Lampronota semirufa Desvignes, 1856 : 81. Holotype <j>, [Great Britain] (BMNH) 
[examined] . 



33o M. G. FITTON 

Desvignes' range of length ('3-4 lines') is presumed to include his var. 1 and therefore the 
only known type-specimen is regarded as holotype. 

Identity. Lissonota semirufa (Desvignes). 

Lampronota varipes Desvignes, 1856 : 81. LECTOTYPE $, [Great Britain] (BMNH), 
here designated [examined]. 

Paralectotype $, same data as lectotype (BMNH). 
The lectotype lacks the tips of the hind tarsi. 
Identity. Lissonota varipes (Desvignes). 

Mesoleptus undecimnotatus Desvignes, 1856 : 33. Holotype $, [Great Britain] (BMNH) 
[examined] . 

The holotype lacks the tips of the antennae and hind tarsi. 

Identity. Junior synonym of Mesoleptidea cingulatits (Gravenhorst) (Morley, 191 ic : 227, 
as Mesoleptus). 

Phytodietus obscurus Desvignes, 1856 : 69. Lectotype $, [Great Britain] (BMNH), 
by fixation of Kerrich (1962 : 51) [examined]. 

Unfortunately the other specimens from Desvignes' type-series cannot be identified with 
certainty. 

Identity. Phytodietus obscurus Desvignes (Kerrich, 1962 : 50). 

Phytodietus ornatus Desvignes, 1856 : 69. Holotype $, [Great Britain] (BMNH) [exam- 
ined] . 

Identity. Phytodietus ornatus Desvignes. 

Pimpla opacellata Desvignes, 1868 : 174. Holotype <j>, [Great Britain] (BMNH) [exam- 
ined]. 

Identity. Junior synonym of Pimpla turionellae (Linnaeus) (Perkins, 1941 : 645). 

Tryphon bimaculatus Desvignes, 1856 : 43. Holotype 6*. [Great Britain] (BMNH) 
[examined] . 

Identity. Juni